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ANNALS OF THE ANNALE VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME, 95 BAND 95

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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 95 BAND

ik VRUSTEES OF THE DIET RRUSREES, VAN GDIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD

1985

EIS ORF CONTENTS

Page
Cooper, M. R.
A revision of the ornithischian dinosaur Kangnasaurus coetzeei Haughton, with a
elassinicanonoLine Ormithischiay (Rublishedtiuned9Ss.) ers eee ee 281
GosLiner, T. M.
The aeolid nudibranch family Aeolidiidae (Gastropoda, Opisthobranchia) from
MmoOpicalsouuneneAuricas (kublishedunel9 85>) many eee eer 233
GRIFFITHS, R. J.
Description of a new South African arminacean and the proposed re-instatement of
the genus Atthila Bergh (Mollusca, Opisthobranchia). (Published June 1985.)... 269
KENNEDY, W. J. & KLINGER, H. C.
Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family
Kossmaticeratdae Spath> 1922° (Publishedtiune 1985.) ey = aes esas ee 165
KENSLEY, B.
The faunal deposits of a Late Pleistocene raised beach at Milnerton, Cape Province,
SoOumpaunicas(aublishedvApnill985s)) 2 te 455s ye eee ee eee oe ltt
KLINGER, H. C. see KENNEDY, W. J.
Orson, Si L.
Early Pliocene Procellariiformes (Aves) from Langebaanweg, south-western Cape
BIOMiNcee SOUUNEAtICaa(eublishedrApmil 1985.) as... one ase ee] ane is coe 123
Otson, S. L.
An early Pliocene marine avifauna from Duinefontein, Cape Province, South Africa.
(LPUULDNS OSG! ZA OTT USCIS) ee aoe ere ern ene eR OE Oo ann Ae) ee esc) Gea 147
SCHOLTZ, A.
The palynology of the upper lacustrine sediments of the Arnot Pipe, Banke,
Namaqualand (rublishedtAvornllOSs:). 55.4545. 5e meen ore ee ie 1

Volume 95 is complete in 8 parts.

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VOLUME 95 PART 1 APRIL 1985 ISSN 0303-2515

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann, Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
April 1985 April
Part 1 Deel

THE PALYNOLOGY OF THE
UPPER LACUSTRINE SEDIMENTS OF THE
ARNOT PIPE, BANKE, NAMAQUALAND

By
A. SCHOLTZ

Cape Town Kaapstad

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THE PALYNOLOGY OF THE UPPER LACUSTRINE SEDIMENTS
OF THE ARNOT PIPE, BANKE, NAMAQUALAND

By
A. SCHOLTZ

Department of Archaeology, University of Stellenbosch
(With 21 figures and 2 tables)

[MS accepted 7 December 1983]

ABSTRACT

The Arnot Pipe on the farm Banke, Namaqualand, is one of the most southerly in the
Gamoep cluster of volcanics, which include ‘kimberlite’ and olivine-melilitite volcanic pipes. The
upper sediments of this pipe, in which palynomorphs as well as plant macrofossils and vertebrate
remains are found, are of lacustrine origin, having been laid down in a small, deep crater lake
formed in a vent of a ‘kimberlite’ volcano. Radiometric dates from other pipes in. the. cluster
indicate that the volcanic activity occurred between 64 and 71 Ma ago. In this study the systematic
palynology of the top 20 m of polleniferous sediments is described.

Seventy-two forms of spores, conifer and angiosperm pollen are described and illustrated
and 64 are formally classified. Fourteen new species and one new genus are defined and
21 further forms are, as far as is known, undescribed in previous literature. The possible affinity
of the fossil forms to 28 plant families, including, amongst the angiosperms, the Proteaceae,
Restionaceae, Ericaceae, Epacridaceae, Euphorbiaceae, Thymelaeaceae, Chloranthaceae,
Casuarinaceae, Cornaceae, Caesalpinaceae and Anacardiaceae, are suggested. The relevance of
these observations to hypotheses about the nature of Palaeogene vegetation in the African
subcontinent are discussed. The Banke palynoflora is contrasted with those of a similar time
range from tropical Africa and Australia to highlight what is unique about the combination of
taxa present in an early Tertiary vegetation in southern Africa. Particular attention is paid to
discussing the early history of some taxa that are at present characteristic elements of the
Capensis Flora.

CONTENTS

PAGE

MITER OCU HOM ae aes eae toi nd CN pe Ny hen gat ale Lahn 2
The location and geology of the Arnot Pipe ................ 3
Previous studies of fossil material from the Arnot Pipe....... 7
DY AGIIN ES Mapp eee eh, cee eect ae ore hig No meetreen U8 ace net eas 9
Waterialtandumethodsieio ae, ane nmeege cides so en ere 10
Rr ANOL OD Weg rere is eA Ce ate See ela, echt ca boe aie eae eat 13
Species list and register of type specimenms.............. 14
IDESCHIPTOMS Aon A Rie y. oN eR ee areal aes Lette cg ae 7)
OME Mi COUMES tvs syaeens nce eesti vet curt gtie eee le ees 88
DISCUSSION set epee eta ht he. eet flies oer ater « tee tare me tt tuegn al eee aw! 88
PNCKNOWICUSEMEMUSi te ie ot ee See e aa coed 103
NCTC TENCE Sr mibiar ney fe ential Rd hc: 5 MRS ky MAN ag ann leh einen 103

1

Ann, S. Afr. Mus. 95 (1), 1985: 1-109, 21 figs, 2 tables.

D ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

This paper presents the first detailed account of an early Tertiary
palynological assemblage from the subcontinent of southern Africa. To date,
direct fossil evidence relating to late Cretaceous and Tertiary vegetation history
and the evolution of plant groups in the region is very limited. The few relevant
palynological studies are mentioned below.

Cursory studies have been published on the Arnot Pipe (Kirchheimer 1934),
on the Knysna lignites—which may relate to a brief period in the Neogene
(Thiergart et al. 1963)—and on a DSDP sequence of mid- to late Cretaceous
marine sediments off the south-western Cape (McLachlan & Pieterse 1978).
Morgan (1978), also studying DSDP material, described late Cretaceous
assemblages from the Angola Basin. A brief study by Scholtz & Deacon (1982) of
sediments from kimberlite pipes in Botswana has provided evidence for the rapid
_ penetration of the ancient austral conifer forest by an early angiosperm flora
during the mid- to late Cretaceous. Sah (1967) has published a detailed
palynological study of the late Neogene sediments from Burundi, just south of the
equator. Finally, Coetzee has done pioneering work on material from numerous
short coastal sequences in the south-western Cape (Coetzee 1978a, 1981). This
work will provide the first detailed palynological evidence about the nature of
vegetation and history of vegetation changes during periods of the Neogene in
this region.

There is, however, a long history of speculative thought on the subject of
vegetation history and the evolution of certain families within the region, based
on analysis of the botanical present (Levyns 1938, 1952, 1964; Adamson 1958;
Taylor 1978; Goldblatt 1978). In the case of the more recent studies their
evidence consists mainly of present patterns of biogeography viewed against the
background of salient facts of plate tectonics. Much better use of the admittedly
patchy African fossil evidence was made by Axelrod & Raven (1978), who
combined fossil evidence, together with the two lines of evidence already
mentioned, to produce their major work on the vegetation history of Africa.
Since the fynbos vegetation of the Cape is a particularly striking component of the
vegetation of southern Africa, being accorded the status of a plant kingdom on its
own (Goldblatt 1978) and possessing a remarkable degree of endemism, and since
the early history of a number of its characteristic taxa appears to be readily
deducible from present patterns of distribution (Levyns 1964), many hypotheses
have been advanced about its origin and evolution.

It was in this context of, on the one hand, an extremely limited fossil record
and, on the other, much interest, many hypotheses and some knowledge of
biogeographical and plant taxonomic patterns relevant to an historical perspec-
tive, that a project to produce detailed palynological evidence on the composition
of vegetation in the southern African subcontinent during earlier phases of
history subsequent to the rise of the angiosperms to dominance of world floras
was undertaken. The sediments of the Arnot Pipe were known to be of late
Cretaceous to Eocene age (Estes 1977) and thus a restudy of this material was

PALYNOLOGY OF THE ARNOT PIPE 3

indicated. The importance of the Banke (Arnot) site had been emphasized by
Goldblatt (1978: 416) and such references to the sketchy evidence previously
available only increased the desirability of a thorough palynological study of the
site.

A preliminary study revealed that careful processing could produce high
pollen concentrations, that the palynomorphs were in a good state of preserva-
tion, and that the pollen assemblage was much more diverse than Kirchheimer’s
work (1934) had indicated.

As part of this project samples of lacustrine sediments were obtained from
numerous other volcanic pipes in the Namaqualand region and from further
afield, and in this paper reference is sometimes made to observations obtained in
the process of ongoing work on this material.

The aim of this study has been to systematically describe the range of forms
found in the top 20 m of polleniferous sediments (52—107 feet (c. 16-33 m) below
surface) of the Arnot Pipe and to optimize the value of these descriptions for
botanists and plant geographers by providing as many pointers to the natural
affinities of the pollen forms as was possible. It must be noted that the latter aim
was achieved almost exclusively by reference to the body of literature available to
the author. This represents an essential aspect of research, but complementary
aspects such as would be provided by the availability of a comprehensive pollen
reference collection of the floras of the subcontinent or published regional pollen
floras did not, in this study, enjoy their rightful place. The level of certainty with
which affinities are suggested is therefore variable, but this should be clear from
the text.

THE LOCATION AND GEOLOGY OF THE ARNOT PIPE

The Gamoep cluster of volcanic pipes associated with olivine-melilitite and
related rocks (Moore 1979) is located in the north-western Cape about 80 km
south of the Orange River and 100 km inland, and is centred around the hamlet
of Gamoep (Fig. 1). The cluster contains upwards of 270 pipe-like bodies
distributed with a north-north-easterly trend. It may be composed of a number of
smaller clusters (Cornelissen & Verwoerd 1975). The cluster straddles the
boundary between the Bushmanland plateau, at an elevation of around 1 200 m,
and the highly dissected escarpment area of Namaqualand. The result is that on
the plateau the pipes are typically buried under metres of sand and exposures are
poor, while in dissected country the volcanic plugs, necks or sediment-filled
depressions are exposed. The Arnot Pipe (30°22’S 18°26’E) on the farm Banke is
one of the more southerly of the pipes in the cluster and is located in a zone of
mildly dissected country between the plateau and the more highly dissected
country of Namaqualand proper. Moore (1979) states that a second distinct
cluster of pipes, associated as far as is known only with olivine-melilitite rocks, is
found between the villages of Garies and Bitterfontein, 50 km south-west of the
Gamoep cluster and closer to the coast.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

67.9m.yr

64.2m.
11.6 mr
|

e?
@e® l 66,7 m.yr

oo A ® puateaxkies
GAs GOREN

\ \

‘N

1A SS
x
e N
Narnor, X
Xx

GARIES \

\ ;
KLIPRAND
i)
. }

A Radiometrically dated volcanics
e Sediment filled pipes
X "Kimbernlite” pipes

. ;
BITTERFONTEIN ©

D Olivine-melilitite pipes

Fig. 1. A map of the Namaqualand—Bushmanland region of the nerth-western Cape showing the

nature and distribution of volcanic pipes in the Gamoep and Garies clusters, and the location of

dated occurrences (after Cornelissen & Verwoerd 1975; Moore 1979). Note: The majority of the
more than 270 pipes in the Gamoep cluster are not shown.

PALYNOLOGY OF THE ARNOT PIPE 5

Rogers (1911), Reuning (1931), and Cornelissen & Verwoerd (1975) have
described three categories of pipes that occur in the Gamoep cluster. Olivine-
melilitite and olivine-nepheline-melilitite pipes often form conspicuous brown,
domed hills in the dissected country. There are a limited number of occurrences
where weathered ‘kimberlite’ or ‘pseudo-kimberlite’ is exposed at the surface, but
the majority of occurrences consist of sediment and breccia-filled diatremes.
These are mostly, if not always, underlain by ‘kimberlite’ and at the present
surface display two modes of crater infilling. The minority are breccia fills in
which the sediment is disturbed by numerous large and small blocks of country
rock testifying to repeated explosive events. The majority are fills consisting of
fine-grained weathered kimberlite, carbonaceous shales and mudstones deposited
under lacustrine conditions. Doubt still remains about the exact mode or modes
of eruption of these ‘kimberlites’, which in this cluster have produced relatively
narrow pipes, some containing in the order of 300 m of bedded lacustrine
sediment. Figure 2B is a geological section through one such sediment-filled pipe,
Koppieskraal K5 (after Cornelissen & Verwoerd 1975).

The Arnot Pipe is an occurrence of the latter type. The diameter of the pipe
is 280—325 m (Reuning 1931) and it is known to contain carbonaceous mudstones
to a depth of at least 135 m (H. Jenner-Clarke, Aram Minerals, pers. comm.).
During the early 1930s a prospecting pit located towards the middle of the pipe
was sunk to a depth of 36 m to investigate the contents of the pipe (Fig. 2A). Ina
paper concerning the composition and geochemistry of the lower section of
sediments exposed in the excavation Reuning (1934) reached two conclusions of
relevance to the present work. Firstly, the sediments of the Arnot Pipe consist of
weathered ‘kimberlite’. This is an important point in suggesting the possible age
of the pipe (see section on ‘dating’). Secondly, since the lower sediments of the
pipe were composed of fine weathered and transported ‘kimberlite’ material with
an almost complete absence of derivatives from the country rock, Reuning
suggested that they were derived from the cone of a strato-type volcano that,
within the catchment area, entirely blanketed the country rock (Namaqualand
gneiss). In an earlier paper, however, Reuning (1931) had suggested another, or
complementary, reason that could explain this phenomenon as well as the lack of
heavier minerals such as ilmenite (derived from the ‘kimberlite’ itself) in the
mudstones of the pipe. A flat landscape together with the luxuriant plant growth
indicated by the rich fossiliferous nature of the sediments could have resulted in a
general low efficiency of water transport and selective deposition of finer
sediments towards the centre of the pipe. This suggestion is supported by the
work of Hawthorne (1975), who confirms that at least in the larger sedimentary
basins of the pipes found in the Botswana Kimberlite Province, coarser material is
selectively concentrated around the margins of the pipes. In the same paper
Hawthorne suggests that the actual volume of material ejected by these types of
volcanoes may have been quite small and that the cones of ejectamenta would
have been correspondingly low.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

main shaft

Be gneiss

mudstone

water table

sandstone
layers - carbonaceous mudstone

dip 30-40
dip 55°

buff coloured fine grained
“kKimberlitic’ sediments interbedded
with carbonaceous mudstone.
rich in macrofossils, frogs. leaves
and branches
fewer frogs, but richer accumulation
of leaves

80 ft

arkose and grit

carbonaceous mudstone

conglomerate

tuffaceous
“kimberlite”

"kimberlite’with
blocks of Country
rock

Fig. 2. A. A stratigraphic section of the known sequence of the Arnot Pipe. The upper 33 m
(107 feet) (maximum depth of the 1929 excavation) at the main shaft and the side excavations are
shown after Reuning (1931). (Reuning’s measurements are given in feet.) The provenance of
samples examined in this study is indicated. The carbonaceous mudstones were recorded to a
depth of 135m in drilling done by H. Jenner-Clarke (pers. comm.) during the 1960s.
B. A stratigraphic section based on the logs of two cores of the Koppieskraal K5 pipe. This can be
taken as a representative reconstruction of the stratigraphy of sediment-filled pipes in the
Gamoep cluster (after Cornelissen & Verwoerd 1975).

PALYNOLOGY OF THE ARNOT PIPE 7

A final point in respect of the origin of the fine sediments of the pipe and the
lack of coarser sediments needs to be made. In Reuning’s (1931, 1934) arguments
he assumed that the substrate in the area at the time of deposition would have
been the country rock, Namaqualand gneisses. However, inclusions of Dwyka
shales have been observed in pipes (Moore 1979: 9) and in the present study an
odd palynomorph specimen of Permian—Triassic provenance was observed. This
indicates that at the time when sediments were being deposited in the crater lake
some Dwyka cover was still present in the area.

From this survey of what is known of the geology of the pipes the following
picture emerges of the local environment during the time of eruption and
sedimentation of the pipes. A great many strato-type ‘kimberlite’ volcanoes
erupted during a relatively short time in a small area. (Using an estimate of a total
of at least 350 pipes in a region of 8 400 km? and the present span of radiometric
dates of 7 Ma, the following calculations can be made: The average density of
pipes is one per 24 km’, though they may be concentrated in subclusters with
densities of about one per 4 km’; average time between eruptions 17 000 years.)
Many small crater lakes could have been synchronously present in the region
although, taking into account the ease with which the ejectamenta could have
weathered, each crater may have had a relatively short life of sediment capture
before infilling was completed.

Using the range of estimates of sedimentation rates for alluvial fan sediments
given by Hooke (1968) and Beaty (1970) it would require between 300 000 and
4 000 000 years to accumulate the possible depth of sediment present in the Arnot
Pipe (135-300 m). The rate of sedimentation, however, would decrease as the
cone height and supply of tefra was reduced.

The basins would presumably have remained as swamp-like features for a
longer period of time and, due to compaction or shrinkage of the initial
sedimentary mass, may have retained a minor ability to capture sediments. The
steeply inward-dipping strata often observed in the pipes (see Fig. 2) is evidence
that this may have occurred. The small drainage basins feeding the lakes would
have been largely unrelated to the developed drainage patterns of the region.

The general landscape was probably dominated by the small emergent cones
of the ‘kimberlite’, strato-type volcanoes in various stages of erosion and, for the
rest, there is little reason to suppose that a relatively flat, mature landscape did
not exist (Mabbutt 1955). Although in the present the peaks of the nearby
Kamiesberge rise some 150-200 m above the plateau, most of the present relief
of Namaqualand is probably the result of more recent incision.

PREVIOUS STUDIES OF FOSSIL MATERIAL
FROM THE ARNOT PIPE

During the 1930s, Reuning and later Boonstra selected samples of material
from the dump of the main excavation. Boonstra (unpublished notes in the South
African Museum) roughly indicated the stratigraphic provenance of a series of

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Material from the 1929 excavation of the Arnot Pipe collected and provenanced by E. Reuning
and L. D. Boonstra and analysed in this study.

Provenance of sample Description of the material

52-58 feet (16-18 m) Bituminous, carbonaceous mudstone.

Unprovenanced Bituminous, carbonaceous mudstone.

65-70 feet (20-21 m) Buff-coloured finely laminated mudstone.

70-90 feet (21-27 m) Buff to grey-coloured mudstone.

90-100 feet (27-30 m) Buff-coloured mudstone with fine silty laminations.

100-107 feet (30-33 m) Buff-coloured mudstone.

100-107 feet (30-33 m) Brown-coloured mudstone with large flecks of organic material.

five small samples (see Fig. 2A and Table 1). A considerable amount of material
was deposited at the South African Museum and the following categories of fossil
material were later studied by various researchers: leaves (Rennie 1931), wood
(Adamson 1931), frogs (Haughton 1931), and palynomorphs (Kirchheimer 1934).

The fossil leaf collection consisted of 70 fragments of dicotyledonous leaves
(and a single fern frond) and Rennie recognized at least twelve different types. In
his short paper only six forms are illustrated by rough line-drawings. The most
common form was described as strap-shaped with serrate margins, and although
Rennie did not do so, it could be described as sclerophyllous. This form was
tentatively compared to the leaves of Myrica, the comparison to some extent
being based on identifications contained in the work of Berry (1925) on Upper
Cretaceous leaf assemblages from North America (Rennie 1931: 252). Rennie’s
tentative comparison is rendered less likely by Chourey’s (1974: 131, 145)
thorough criticism of Berry’s work. Chourey noted that in the early years of study
of leaf fossils of late Cretaceous—Tertiary age, inadequate identification criteria
were employed and many identifications made then can no longer be accepted. In ~
particular many fossils were identified either as myricaceous or proteaceous;
affinity with the genus Banksia of the Proteaceae was regularly suggested. In fact
this tentative alternative identification was made by Berry in the very paper cited
by Rennie. It should be noted that, while rejecting most of the Myrica identifi-
cations made on late Cretaceous—early Tertiary material from North America and
Europe, Chourey (1974) suggests that the phenomenon of a world-wide occurrence
and prominence of this form type is significant and worthy of further study. The
‘myricaceous’ leaves from Arnot are part of this world-wide phenomenon.

It was not possible to obtain adequate descriptions of the remaining eleven
leaf types to allow for identification and Rennie merely noted their general
‘mesophytic’ habit.

Adamson (1931) studied silicified wood samples from opalized sections of the
superficial sandstone layers found close to the contact between the pipe fill and
the gneiss. He identified the fossil wood as that of Ficus cordata, which grows in
the area at present. Kirchheimer (1934: 47) quotes Reuning’s statement that these
sandstone layers may be of a substantially younger age than the underlying clays.

PALYNOLOGY OF THE ARNOT PIPE 9

Haughton (1931), in the most detailed of these early studies, described a
single new species of Pipidae, Eoxenopoides reuningii and, in terms of relatively
conservative pipid evolution and lacunae in their fossil record, could only assign a
Cretaceous to early Tertiary age to the form. Estes (1977) agreed with
Haughton’s systematic description and on morphological grounds also upheld
Haughton’s age bracketing. However, on extraneous grounds he favoured an
Oligocene age for the sediments (Estes 1977: 51).

Kirchheimer (1934) cursorily described six palynomorph types including two
disaccate and four triaperturate angiospermous forms. He also mentioned that
spores and inaperturate grains had been observed. He obtained his best results
from thin sectioning of opal concretions and obviously encountered difficulties in
processing the carbonaceous clays of the lower layers available to him.

DATING

The best biostratigraphic evidence on which to base an age estimate for the
sediments of the Arnot Pipe has been produced by the present study and is
discussed in a later section in the context of the radiometric evidence mentioned
below. The suggestion by Axelrod & Raven (1978), based on an extremely
tenuous comparison with the North American and Mediterranean plant macro-
fossil record, that the occurrence of a sclerophyllous leaf type indicates a late
Eocene to Miocene age, can be regarded as insubstantial. The arguments
advanced by Reuning (1931) and Haughton (1931) on sedimentological and
geomorphological grounds for a late Cretaceous age were probably never
intended to be more than speculative.

The most positive dating evidence relevant to suggesting a possible age for
the Arnot sediments consists of a number of radiometric determinations on
material from pipes in the Gamoep cluster (Davis 1977). The dates and location
of pipes are given in Figure 1. All the dates were obtained by the *°U/*"°Pb
method applied to zircon inclusions in kimberlitic material. Until these dates were
available, the single K/Ar date of 38,5 Ma from an olivine-melilitite pipe on the
farm Dikdoorn in the Garies—Bitterfontein cluster was the most pertinent
radiometric determination relevant to the possible age of the Gamoep cluster.
This was also the only date available when the dating of the Arnot sediments was
last discussed (Estes 1977).

In view of the range of dates presently available, the evidence is strongly in
favour of accepting a 60-70 Ma age bracket for volcanic activity in the Gamoep
cluster and for the Arnot Pipe. Firstly, all three pipes dated in the Gamoep cluster
are apparently ‘kimberlitic’, as is the Arnot Pipe. Secondly, the five dates
available are relatively tightly grouped within a 7 Ma time span and the Arnot
Pipe is within 30 km of the dated pipes. No olivine-melilitite pipes from the area
have yet been dated.

Two K/Ar dates from olivine-melilitite pipes in the small cluster south of
Garies are available and are younger or considerably younger than the dates for

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

the main Gamoep cluster, i.e. 54,1 and 38,5 Ma. It must be pointed out that the
younger date here and the other two Oligocene dates shown in Figure 1 have in
fact not been fully published (Kréner 1973) and should be treated with caution. In
contrast to the use of the *°U/*%°Pb method to date the pipes in the Gamoep
cluster, the K/Ar method was used in the case of the Garies—Bitterfontein pipes.
Excluding possible problems involved in the dating techniques, the spread of
dates from the Garies—Bitterfontein pipes as well as from olivine-melilitite pipes
wider afield (see Fig. 1) may indicate that the time span of volcanic activity either
in this small cluster or throughout the distribution of olivine-melilitites was much
greater than that of the volcanic activity associated with the ‘kimberlite’ pipes of
the Gamoep cluster. Moore (1979) (see below) has suggested at least two
mechanisms that could explain the phasing of volcanic activity resulting in
extrusion of lighter ‘kimberlite’ earlier than that of more dense olivine-melilitite.
He (1979: 136) also discusses some disparities in results obtained in the
radiometric dating of kimberlites, olivine-melilitites and related rocks, and
suggests that many determinations are possibly questionable.

Moore (1973, 1979) develops hypotheses that might explain:

(a) the possible younger ages of the olivine-melilitite versus the ‘kimberlite’
volcanics of the region;

(6) the possible pattern of younger dates occurring closer to the coast; and

(c) the possible concomitant pattern of progressive increase in the magnesium
oxide content of volcanics from the olivine-melilitites of the Garies—

Bitterfontein cluster through those of the Gamoep cluster to the ‘kimberlites’

of the latter region.

These hypotheses involve the late Cretaceous—Oligocene epeirogenic uplift
along the warp axis of the western escarpment as a primary cause of volcanic
activity. Furthermore, either the fractionation of the parent magma in the stress
zone beneath the warp and a resultant enrichment of a less dense portion with
volatiles could lead to a first phase of volcanic activity in which ‘kimberlite’ was
extruded, or the progressive thickening of the craton towards the interior of the
continent might cause extrusion of magmas from different depths and thus
explain the coast—interior gradient of geochemical attributes and perhaps dates.

So much for the possible dating of the volcanics that provides a maximum age
for the lacustrine deposits. As suggested it is likely that infilling of the craters
would have proceeded relatively rapidly, or would at least at first have been
rapid. The estimates obtained from sedimentological work in similar environ-
ments suggest that a maximum of 4 Ma could have been required to accumulate
the depth of deposit that may be present in the Arnot Pipe.

MATERIAL AND METHODS

As already stated, the material curated in the South African Museum was
collected by E. Reuning and L. D. Boonstra during the 1930s. Even the five small
samples whose stratigraphic provenance was indicated by Boonstra (see Fig. 2A

PALYNOLOGY OF THE ARNOT PIPE 1]

and Table 1) were probably collected from the dump (which was presumably to
some extent systematically organized) after the excavation had been closed. This
is inferred from the fact that this small series includes samples from the lowest
levels reached in the excavation, yet the work had ceased at this level (35 m) due
to the instreaming of water, and the pit would presumably have filled with water
to the level of the water-table 20 m higher up. The coherent pattern of the pollen
diagram (Fig. 3) does to some extent suggest that the provenancing of the samples
is correct.

The seven samples studied are listed and described in Table 1, in the same
sequence as their pollen spectra appear in the pollen diagram.

The bulk of the collection was unprovenanced, but since this material often
contains macroscopic fossil material, it is unlikely to have come from the upper
third of the excavation. Reuning (1931) noted that preservation of fossil material
above the level of the water-table was poor. On the other hand, in view of the
pollen counts (Fig. 3), it seems unlikely that the bulk of the samples could have
come from below the 18 m (60 foot) level. The combined count of two counts on
the unprovenanced material is placed below those of Boonstra’s 52-58 foot
(16-18 m) samples on the pollen diagram and are virtually indistinguishable from
the latter.

The unprovenanced material is of two sedimentological types—a dark,
carbonaceous mudstone and a buff-coloured mudstone. The pollen spectra from
the two types are similar and unfortunately, since the best-preserved and richest
concentrations were obtained from this material, many holotypes designated in
this study are located in preparations from it.

It appeared that two sediment types also occurred between 30 and 33 m
(100-107 feet), but no indication was given about their stratigraphic relationship.
In Table 1 and Figure 3, the spectrum from the buff-coloured mudstone 1s
arbitrarily placed above that of the dark, carbonaceous mudstone.

Kirchheimer used a single-stage process, cold hydrofluoric acid digestion, to
concentrate palynomorphs. The stratigraphic provenancing of his samples was
uncertain. In the present study a six-stage process was employed, consisting of the
following steps:

1. Crushing and dispersal of the sample in a 0,3M solution of tetrasodium
pyrophosphate, followed by numerous short centrifuges and rinses in deionized
water to wash out the fine clay fraction. If necessary, additional applications of
tetrasodium pyrophosphate solution were used. This treatment is an adaptation
of the process described by Bates et al. (1978) but was independently suggested by
a soil scientist, J. J. N. Lambrechts of the Department of Agriculture, University
of Stellenbosch.

2. Standard zinc chloride heavy-liquid flotation to separate organic and
larger-sized inorganic fractions.

3. In the treatment of a few of the more heavily carbonaceous samples it was
necessary to cause some oxidation using either 30 per cent hydrogen peroxide
solution or nitric acid.

[2 ANNALS OF THE SOUTH AFRICAN MUSEUM

eS 3s ¢ & SSeS a ee Ss
, 1 = © es ) w o co a 8 4! > ~
as" 38 Ss 2 8 4 via: eo en tee PROVENANCE OF SAMPLE
— ~ . - —_- = oe o Qo mm [.s}
PS = — a — a = ] e - =< e@
= = = = =
oO = = oO
= =
STEREISPORITES TRIORITES OPERCULATUS
OTHER SPORES
| J DICOLPOPOLLIS
TRICOLPITES RETICULATUS
CLAVATIPOLLENITES
TRIORITES SPHERICUS
PROPYLIPOLLIS
ERICIPITES
PODOCARPIDITES
NILFOROIA
“ LE he 8 ARAUCARIACITES
Pd
™m
TRICOLPOROPOLLNITES ARNOTIENSIS
uw
cy
[=]
se
E a R i R ry Wm TRICOLPOROPOLLENITES BRINKIAE
S233 S & S EB poruen sun GT OW TM ™ rerpororerravires spuerrcus

Fig. 3. Pollen diagram showing relative abundance of those taxa that constitute greater than
1 per cent of the grains present at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 1

4. Standard acetolysis process.

5. A two or three-minute rinse in hydrofluoric acid to destroy any remaining
silica particles.

6. A rinse in warm 10 per cent hydrochloric acid.

After final washing with deionized water the concentrates were suspended in
a 50 per cent glycerol solution for light microscopy or left in deionized water for
mounting on SEM stubs. Permanent slides were made with glycerine jelly and
sealed with nail varnish.

The full series of slides with holotypes and paratypes ringed and documented
is deposited in the South African Museum. A register of South African Museum
catalogue numbers for these holotypes and paratypes is given in the species list
(e.g. SAM-K6155). A duplicate set of slides and the sealed phials containing the
remaining concentrates is deposited in the Department of Archaeology,
University of Stellenbosch. Counting of samples and photomicroscopy was done
on a Wild M11, and SEM work was done on a JEOL JSM-35 housed in the
Department of Physics, University of Stellenbosch.

PALYNOLOGY

Kemp & Harris (1977: 5) provide a recent review of the problems
encountered in the systematic palynology of early Tertiary material. They state
that “Tertiary palynology even more than the palynology of older sedimentary
rocks, has suffered from a marked ambiguity of approach’. This ambiguity of
approach arises in that three different approaches to nomenclature have been
applied. Authors have variously assigned fossil palynomorphs to extant genera,
used a name that suggests affinity to an extant genus (e.g. Araucariacites for forms
resembling pollen of the genus Araucaria), or applied an artificial name based on
morphological criteria alone (e.g. Triorites or Monocolpopollenites).

In addition, a certain regionality of nomenclature, reflecting the isolation in
which early palynological work was done, is inherited by present-day palynol-
ogists (depending on what literature they are exposed to). This problem can be
compounded by the difficulties entailed in keeping up to date with more recent
work published in an array of journals.

Most palynologists working on early Tertiary material have elected, firstly, to
continue to use the binomial system of nomenclature together with the standard
botanical rules of typification, priority, etc. This promotes stability and some
measure of uniformity in the use and creation of names for fossil palynomorphs,
and is the system adhered to in the present study. However, it does not
discourage the proliferations of names and the growth of ‘portmanteau’ genera.
Secondly, in establishing new genera palynologists have favoured the artificial
system of nomenclature, which promotes the utilization of palynology as a strati-
graphic tool (Sah 1967: 6), and this practice is also followed in the present work.

In this study the descriptive and stratigraphic palynological literature from
Australia, India and tropical Africa has been most often consulted. As could be

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

expected, in view of its isolation from these areas, the palynomorph assemblage
from Banke appeared unusual and many new specific names have resulted. Both
in the occurrence of a number of unique forms and in the general composition of
the assemblage, the already developed distinctiveness of the flora of the
subcontinent is apparent.

A new specific or generic name was not proposed unless the form concerned
was reasonably common in the Arnot samples. A large number of forms were
present at low frequencies and in these cases, if at least three specimens in good
state of preservation were observed, the forms were described and illustrated,
placed if possible in a genus, and their affinities suggested. This was deemed to be
worthwhile in terms of the aims of the study and the unique nature of these
observations at present. Other forms will no doubt be systematically described as
work on the pollen and spore assemblages from ‘“kimberlite’ pipe occurrences in
the northern Cape and Botswana is extended. Sequences are already known in
which elements rare at Arnot are common or even dominant. One form, referred
to informally as Fenestriorites, is not described in the present paper as it is being
described elsewhere. The form promises to be an especially important marker
species in the regional sequence. Some importance is therefore placed on its
occurrences in the lowest levels as yet sampled at Arnot (where it is extremely
rare) and this is discussed in the section on interpretation of the present
palynological evidence.

SPECIES LIST AND REGISTER OF TYPE SPECIMENS
Spores

Trilete spores
Stereisporites sp.
Cyathidites australis Couper, 1953
Planisporites sp.
Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968
Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973
Foraminisporis sp.
Herkosporites elliottii Stover, in Stover & Partridge, 1973
Camarazonosporites bankiensis sp. nov.
Holotype SAM-K6155
Polypodiaceoisporites sp.
Trilites sp.

Monolete spores
Microfoveolatosporis fromensis (Cookson) Harris, 1965
Cicatricososporites sp.

Alete spores
Reticulatasporites grandis sp. nov.
Holotype SAM-K6156; paratypes SAM-—K6157, K6158

PALEY NOLOGY OFSHHE ARNOT PIPE

Spores not assigned to genus
Forma A
Forma B
Forma C
Forma D

Pollen of Coniferae

Inaperturate pollen
Araucariacites australis Cookson ex Couper, 1953
Araucariacites sp.

Monosaccate pollen
Zonalapollenites sp. A
Zonalapollenites sp. B

Disaccate pollen
Lygistepollenites sp.
Podocarpidites sp.
Podocarpidites kamiesbergensis sp. nov.
Holotype SAM-K6159; paratypes SAM—K6160, K6161, K6162
Podocarpidites riembreekensis sp. nov.
Holotype SAM-K6163; paratypes SAM-—K6164, K6165, K6166

Pollen of Angiospermae

Monoaperturate pollen

—Monocolpate pollen
Arecipites plectilimuratus Chmura, 1973
Arecipites sp. A

Arecipites sp. B
Liliacidites sp. A
Liliacidites sp. B
Clavatipollenites sp. A
Clavatipollenites sp. B
Clavatipollenites sp. C
Monocolpopollenites sp. A
Monocolpopollenites sp. B

—Monoporate pollen

Milfordia hypolaenoides Erdtman, 1960
Milfordia sp.

—Dicolpate pollen

Dicolpopollis sp.

—Monocolpate pollen not assigned to genus
Forma E

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Triaperturate pollen

—Triporate pollen
Triorites operculatus sp. nov.

Holotype SAM-—K6167; paratypes SAM—K6168, K6169, K6170
Triorites sphericus sp. nov.

Holotype SAM-K6171; paratypes SAM—K6172, K6173
Triorites harrisii Couper, 1960
Triporopollenites namaquensis sp. nov.

Holotype SAM-—K6175; paratypes SAM-—K6176, K6177, K6178
Proteacidites sp. A
Proteacidites sp. B
Propylipollis meyeri sp. nov.

Holotype SAM-K6179; paratypes SAM-—K6180, K6181, K6204
Propylipollis sp.
Fenestriorites sp. (not described in this paper)

—Tricolpate pollen

Tricolpites reticulatus Cookson, 1947
Tricolpites sp. A
Tricolpites sp. B
Tricolpites sp. C
Tricolpites sp. D
Crototricolpites densus Salard-Cheboldaeff, 1978
Spinitricolpites jennerclarkei gen. et sp. nov.
Holotype SAM-K6182; paratypes SAM—K6183, K6184, K6185

—Tricolporate pollen

Tricolporopollenites grandis sp. nov.

Holotype SAM-K6186; paratypes SAM-K6187, K6188
Tricolporopollenites arnotiensis sp. nov.

Holotype SAM-K6189; paratypes SAM—K6190, K6191
Tricolporopollenites brinkiae sp. nov.

Holotype SAM-K6193; paratypes SAM—K6194, K6195
Tricolporopollenites coetzeeae sp. nov.

Holotype SAM-K6196; paratypes SAM-K6192, K6174
Tricolporopollenites sp. A
Tricolporopollenites sp. B
Tricolporopollenites spp. C & D

Pollen with more than three apertures

Retistephanocolpites sp.
Grootipollis reuningli sp. nov.

Holotype SAM-K6197; paratypes SAM-—K6202, K6203
Ulmipollenites sp.

PALYNOLOGY OF THE ARNOT PIPE 17

Inaperturate pollen
Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979

Pollen found in obligate tetrads
Ericipites sp. A
Ericipites sp. B
Dicotetradites sp.
Triporotetradites sphericus sp. nov.
Holotype SAM-—K6198; paratypes SAM-—K6199, K6200, K6201

Dicotyledononous pollens not assigned to genus
Forma F
Forma G
Forma H
Forma I

DESCRIPTIONS
Spores

‘The identification of Tertiary spores is more problematical than that of fossil
pollen grains. This is partly because they are less well known but mainly because,
as Knox (1935) and Selling (1946) have shown, a particular type is not necessarily
restricted to a single genus or even family and considerable variation often exists
within a genus’— Cookson (1947: 135).

Trilete spores
Genus Stereisporites Pflug, in Thomson & Pflug, 1953

For a discussion of this genus see Dettmann (1963: 25).

Stereisporites sp.
Fig. 4A—D
Compare

Sphagnum antiquasporites Wilson & Webster, 1946: 273 (fig. 2).

Triletes australis Cookson, 1947: 136, pl. 15 (figs 58-59).

Sphagnites australis (Cookson) Balme, 1957: 15, pl. 1 (figs 1-3).

Stereisporites antiquasporites (Wilson & Webster) Dettmann, 1963: 25,
figs 20-21. Harris 1974: 79, pl. 24 (fig. 20).

Description

Microspore trilete, biconvex, amb subtriangular to subspherical with convex
sides and broadly rounded angles. Laesurae straight and simple, length one-half
spore radius. The exine is uniformly thick and the distal and probably the
proximal surfaces are covered by very low angular rugulae. Equatorial diameter
24-35 pw, exine 1-2 wm.

18

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. A-D. Stereisporites sp. EE. Cyathidites australis. _F—G. Planisporites sp.
H. Foveotriletes margaritae. \—J. Foveotriletes lacunosus. K—L. Foraminisporis sp.

PALYNOLOGY OF THE ARNOT PIPE 19

Remarks

The slight thickenings in the radial regions at the equator and the low, distal
polar thickening, circular in outline, mentioned by Dettman (1963: 25) were not
observed, nor do they appear in either the descriptions or photomicrographs of
Wilson & Webster (1946), Cookson (1947), Balme (1957) or Harris (1974). The
differences between the present form and the species Stereisporites antiqua-
sporites (Wilson & Webster) Dettmann, 1963, are therefore slight, but sufficient
to prevent identification.

Affinity

The genus Sphagnites Cookson, 1953, was established to include fossil spores
resembling those of the peat-moss family Sphagnaceae. (See discussion of
Stereisporites in Boros & Jarai-Komlddi 1975: 9.)

Distribution

The genus is known from Jurassic to Tertiary sediments and is sometimes
common in Australian late Cretaceous sediments, particularly in highly carbon-
aceous samples (Dettmann 1963). Stereisporites sp. is the most common spore at
Banke. Due to the long time range shown by the genus Stereisporites, it may have
little stratigraphic value, but Balme (1957: 15) remarks that it is rarely seen in
marine and transitional sediments and may therefore be important in facies
studies. A Sphagnites form is illustrated but not described from the Knysna
lignites (Neogene?) by Thiergart et al. (1963, table 2, fig. 6) and was also recorded
in sediments of middle to late Cretaceous age off the south-western Cape
(McLachlan & Pieterse 1978). A survey of the literature suggests that Sphagnites
forms are not known from late Cretaceous or Tertiary sediments of tropical
Africa. The modern family is strongly circumboreal in distribution, although one
form is cosmopolitan and other forms occur in the Southern Hemisphere, mainly
in New Zealand (Boros & Jarai-Komlddi 1975). A few Sphagnum species are
found in southern Africa and are common although ‘confined to shaded mountain
seeps, streambanks or swampy areas’ (Magill 1981: 23).

Genus Cyathidites Couper, 1953
See discussion in Dettmann (1963: 22).

Cyathidites australis Couper, 1953

Fig. 4E
Cyathidites australis Couper, 1953: 27, pl. 2 (fig. 11).

Description

The spores are trilete. The laesurae are ciearly defined (about two-thirds of
the radius of the spore), narrow and straight. The ends of the laesurae may

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

terminate in a short bifurcation. The spores are triangular with rounded apices
and the sides are mostly concave in polar view. The exine is thin and psilate, and
both proximal and distal surfaces are convexly curved. Equatorial diameter
46-60 pw, exine 1 p.

Affinity

Couper (1953: 27), having compared the present form to the spores of the
extant fern Thyrsopteris elegans, quoted Copeland (1947: 48) on the latter
species: ‘It may well be a relict from the time when Dicksonia and Cyathea had a
common ancestor.’ On the basis of the available evidence Couper suggested that
Cyathidites australis may be the spore of a tree-fern.

Distribution

Cyathidites australis is widely distributed in the Mesozoic and Tertiary of the
Northern Hemisphere and Australia, and is often abundant. It is a rare
component of the Banke spore flora. Similar forms are present in the late
Cretaceous (McLachlan & Pieterse 1978, pl. 1 (figs 1-2)) and Neogene (?)
(Thiergart et al. 1963, pl. 2 (figs 12, 14—15)) of the southern African subcontinent.

Genus Planisporites Knox, 1950
Planisporites sp.
Fig. 4F-—G

Description

The spore is trilete and its amb is subtriangular to deltoid with rounded
corners. The exine is relatively thick. The distal face and equatorial regions are
decorated with microconi, which are mostly solitary, but may also be arranged in
rows. The bases of adjacent coni may be linked by fine ridges. The proximal face
is psilate and the laesurae are long, thin and simple and almost reach the equator.
The equatorial diameter of the illustrated specimen is 33 w and the exine is 2 wu
thick.

Affinity
There is no information regarding possible affinities of Planisporites sp.

Distribution

Planisporites sp. is rare at Arnot. There is no further information on the
distribution of the genus.

PALYNOLOGY OF THE ARNOT PIPE 2A

Genus Foveotriletes Potonié, 1956
Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968
Fig 4H

Triletes margaritae van der Hammen, 1954: 102, pl. 17.
Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968: 286, pl. 1 (figs 1-2).

Description

The spore is trilete with a roundly triangular amb and is circular to biconvex
in lateral view. The laesurae are straight with finely serrate margins and are one-
half the spore radius. The exine is relatively thin and the whole surface is densely
covered by scrobuli; approximately sixty scrobuli per 100 «*. Equatorial diameter
is 55 w; laesurae 15-18 w; exine 1-2 wp.

Affinity

Spores that resemble F. margaritae are produced by members of the
Ophioglossaceae, especially the genera Botrychium and Ophioglossum (Salard-
Cheboldaeff 1981).

Distribution

Foveotriletes margaritae is recorded from sediments of Palaeocene age in
tropical Africa and South America, where it becomes extinct during the lower
Eocene (Germeraad ef al. 1968). It is rare at Arnot.

Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973
Fig. 4I-J
Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973: 248, pl. 14 (fig. 6).

Description

The spore is trilete and the amb is rounded triangular to subcircular. The
distal surface is convex and the proximal is pyramidal. The laesurae are two-thirds
to three-quarters spore radius, irregular, not straight, and have thin, steep lips.
The proximal surface is psilate, the distal surface is covered with poorly
delimited, shallow foveola that almost encroach on to the proximal face. The
exine is relatively thick, approximately 2 uw, and its inner surface appears to
follow the undulations of its outer surface. Equatorial diameter 35-37 wp.

Affinity
There is no information on the affinity of F. lacunosus.

Distribution

Foveotriletes lacunosus is known from the Oligocene to Miocene in
Australia; it is rare at Arnot.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Foraminisporis Krutzsch, 1959

See discussion in Dettmann (1963: 71).

Foraminisporis sp.
Fig. 4K—L
Compare
Foraminisporis dailyi Dettmann, 1963: 72, pl. 14 (figs 15-18).

Description

The spore is trilete. The amb is rounded triangular to subcircular with a
notch in what is probably a narrow sculptured cingulum (see Dettman 1963: 71)
where the laesurae meet the equator. The outline of the grain is irregular. The
laesurae are straight and placed on sculptured ridges running the length of the
radial areas. The distal face is covered by verrucate to spinulate structures whose
bases sometimes coalesce. The proximal face is distinctly less verrucate with
occasional foveola. Equatorial diameter 45-50 p, cingulum 3 wp.

Affinity
The affinity of the genus Foraminisporis is perhaps with the bryophyte family
Anthocerotaceae (Dettmann 1963: 71).

Distribution

Foraminisporis 1s world-wide in the Cretaceous; F. dailyi is present in late
_ Cretaceous sediments off the south-western Cape (McLachlan & Pieterse 1978).
Foraminisporis sp. was common at Arnot.

Genus Herkosporites Stover, in Stover & Partridge, 1973
See discussion in Stover & Partridge (1973: 248).

Herkosporites elliottii Stover, in Stover & Partridge, 1973
Fig. 5A—B
Herkosporites elliottii Stover, in Stover & Partridge, 1973: 248, pl. 13 (fig. 7).

Description

The spore is trilete, the amb roundly triangular and the laesurae extend
almost to the equatorial margins. The radial area immediately bordering the lips
of the laesurae appears smooth, but in the remainder of the radial region
regularly spaced, thin structures (? folds) arranged at right angles to the laesurae
are present. The proximal interradial area is psilate. The laesurae are narrow with
thin raised lips. The distal surface is spinate; spines are of a uniform height 3-4 yu
with abruptly broadening bases, which sometimes coalesce in fine ridges; space
between spines 1,5—2,5 uw, exine 1 w, equatorial diameter 40-45 w.

PALYNOLOGY OF THE ARNOT PIPE 23

Affinity

There is no direct information on the affinities of Herkosporites. Dettmann
(1963: 36) quotes Cookson & Dettmann (1958) on a comparison between the
morphologically related genus Ceratosporites and certain members of the extant
genus Selaginella.

9046

Fig. 5. A-B. Herkosporites elliottii. C-D. Camarazonosporites bankiensis sp. nov.
E-F. Polypodiaceoisporites sp.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Herkosporites elliottii is distributed in the Palaeocene to Miocene in
Australia; it was rare at Arnot.

Genus Camarazonosporites Pant ex Potonié, 1956
Camarazonosporites bankiensis sp. nov.
Fig. SC-D

Etymology

This species is named after the farm Banke, near Platbakkies, Namaqualand,
and the site name.

Description

The spores are trilete and cingulate with a convexly triangular amb and
rounded apices; biconvex in lateral view. The proximal face is subpsilate,
probably finely granulate. The distal face is covered by a fine hamulate
sculpturing, which is completely lost in the radial equatorial region, following the
characteristic trend for the reduction of the exine of this region in the genus
Camarazonosporites. The narrow laesurae extend to the equator and are
bordered by thin, steep membraneous folds, which thus form irregular lips. The
lips may overfold the laesurae. The cingulum (equatorial crassitude) is 6-7 pu
wide in the interradial regions, and narrows to 1-1,5 w in the radial region, giving
the grain a rounded triangular aspect in plan view. Equatorial diameter is
59-63 p.

Remarks

Camarazonosporites bankiensis is quite common in the Banke material and
can therefore serve as a type population. The spores are twice the size of the type
species of the genus, and much larger than any other known species of
Camarazonosporites. It is also larger than most species in morphologically related
genera such as Coronatispora, Sestrosporites and Camarazonotriletes.

Affinity
The affinities of C. bankiensis are not known. It has a general similarity to
the spores of some members of the Lycopodiaceae.

Distribution

Camarazonosporites bankiensis is common at Arnot. The genus is known
from the late Cretaceous. An apparently similar form, labelled Lycopodium-
type, is present in the Knysna lignites (Neogene?) (Thiergart ef al. 1963, pl. 2 (figs
1—3)) but no description is provided.

PAL Y NOLOGY OPMHE ARNOT PIPE 25

Genus Polypodiaceoisporites Potonié, 1951 ex Potonié, 1956
Polypodiaceoisporites sp.
Fig. SE-F

Description

A trilete, cingulate spore. The amb is triangular with broadly rounded
apices. The laesurae are straight and reach the cingulum without extending into
it. A concavely triangular area in the central radial area of the proximal face is
markedly depressed below a surrounding ridge, which is decorated with robust
rugulate structures. This sculpturing becomes reduced as the laesurae are
approached within the centrally depressed area. The rugulate sculpturing on the
distal face is formed by much flatter and broader structures. The cingulum is
smooth and of variable width. Equatorial diameter 80 w, cingulum 6 w wide.

Affinity
The generic designation suggests affinity to the Polypodiaceae, but the spores
of the genus Pteris of the Pteridaceae (Muller 1968) also resemble this form.

Distribution

The genus Polypodiaceoisporites is known from the Tertiary and Cretaceous
of both the Southern and Northern hemispheres. Polypodiaceoisporites sp. is rare
at Arnot.

Genus Trilites Cookson ex Couper, 1953
Trilites sp.
Fig. 6A-B
Compare

Trilites ohaiensis Couper, 1953 in Couper, 1960: 41, pl. 2 (figs 7-8).
Latrobosporites crassus Harris, 1965: 81, pl. 25 (figs 8-9).

Description

The spore is large and trilete with a relatively thick psilate exine. It is
uniformly covered by a thin outer membrane, which closely adheres to the exine
and is thrown up into low folds and wrinkles to form a dense hamulate pattern.
The laesurae are long and thin and outlined by folds in the perinous membrane.
The laesurae reach or almost reach the equator. It appears that the perinous
membrane withdraws from a small area around the apices of the amb where the
laesurae meet at the equator. This suggests that the grain is limbate. The amb of
the grain is that of a broadly rounded triangle. Equatorial diameter 60 w.

26

Fig. 6.

ANNALS OF THE SOUTH AFRICAN MUSEUM

i ‘ 1 n 40 se

A-B. Trilites sp. C. Microfoveolatosporis fromensis. D. Cicatricoso-
sporites sp. E-F. Reticulatasporites grandis sp. nov.

PALYNOLOGY OF THE ARNOT PIPE 27

Remarks

Despite Harris’s (1965: 81) comment that no forms resembling the new genus
and species Latrobosporites crassus Harris, 1965, were then known from Austral-
asia, there seems to be some measure of resemblance between TJrilites ohaiensis
Couper, 1953 and L. crassus Harris, 1965. There is a resemblance between these
two species and the present form, Trilites sp., although the amb of the latter is
more triangular and neither of the former two species are described as perinate.
Trilites ohaiensis was originally described (Couper 1953: 30) as having a
verrucate—granular sculpture, but this was later revised (Couper 1960: 41) and the
grains described as having a rugulate—vermiculate sculpture. Harris (1965)
described the sculpture of L. crassus as consisting of low interlocking rugulae and
lumina of similar size and shape.

Affinity
Harris (1965) suggested that L. crassus had affinity to the extant Selaginella
cathedrifolia-group as defined by Knox (1950).

Distribution
Trilites sp. is rare at Arnot. Trilites ohaiensis is rare in New Zealand late

Cretaceous sediments (Couper 1960), and L. crassus is common in Palaeocene
sediments from south-western Australia (Harris 1965).

Monolete spores
Genus Microfoveolatosporis Krutzsch, 1959
Microfoveolatosporis fromensis (Cookson) Harris, 1965
Fig. 6C

Schizaea fromensis Cookson, 1956: 43, pl. 8 (fig. 3).
Microfoveolatosporis fromensis (Cookson) Harris, 1965: 84, pl. 24 (fig. 7).

Description

The spores are monolete and oval to circular in polar view and reniform
(concavo-convex) in lateral view. The laesura has distinctly raised lips for most of
its length and is about one-half the total length of the spore. The exine is thick and
robust and is uniformly and densely covered by regularly arranged, shallow
microfoveola. Shallow furrows may connect adjacent microfoveola. The overall
dimensions are very regular, approximately 80 x 60 w; length of laesura 40 yw,
exine 3 w, depth of foveola 0,5 uw, 11-16 foveola per 100 pw’.

Remarks

Harris (1965) does not make it clear how Microfoveolatosporis fromensis
(Cookson) Harris, 1965 differs from Microfoveolatosporis pseudodentatus
Krutzsch, 1959, the type species of the genus.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

The genus Microfoveolatosporis has affinities with the genus Schizaea of the
Schizaeaceae. Spores of Schizaea pennula Swartz, an extant Columbian species,
are indistinguishable from M. fromensis (Murillo & Bless 1978: 356).

Distribution

Microfoveolatosporis is known from the Tertiary of both hemispheres. The
extant genus Schizaea is distributed predominantly in the Southern Hemisphere,
with two species occurring in southern Africa (Welman 1970). Microfoveolato-
sporis fromensis exits from the tropical African record at the Cretaceous—Tertiary
boundary (Salard-Cheboldaeff 1979).

Genus Cicatricososporites Pflug & Thomson,
in Thomson & Pflug, 1953

See discussion of Cicatricososporites and Schizaeoisporites in Jansonius &
Hills (1976: 468-469, 2530) and Srivastava (1971: 256).

Cicatricososporites sp.
Fig. 6D
Compare

Cicatricososporites norissii Srivastava, 1971: 257, pl. 1 (figs 5-8).

Description

The spore is alete or possibly monolete, with canaliculate to cicatricose
sculpturing. Odd ridges may bifurcate for a short distance and the sculpturing is
somewhat asymmetrical, resulting in a slight spiralling appearance. No laesura
was observed, but it may lie parallel to, and be almost indistinguishable from, the
grooves of the sculpturing. The length of the specimen illustrated is 60 pw.

Affinity

The genus Cicatricososporites has affinities with the genus Schizaea.
Srivastava (1971) stated that ‘Cicatricososporites norissii is comparable with
spores of the extant species Schizaea laevigata illustrated by Selling 1946’.

Distribution

Cicatricososporites norissii is common in the lower member of the Edmonton
Formation (Maastrichtian), Canada, where it is a prominent component of a flora
that Srivastava (1971) suggested grew under humid conditions. The distribution
of Cicatricososporites in the Palaeocene presents a strange disjunct pattern, which
is reflected in the present-day distribution of the genus Schizaea (see Table 2).
Only one specimen of Cicatricososporites sp. was observed at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 29

Alete spores
Genus Reticulatasporites Ibrahim, 1933

Ibrahim’s (1933) diagnosis of the genus Reticulatasporites was very general:
‘Spores without trilete mark, and with a measurable reticulate sculpture on the
spore wall; meshes up to [or as small as ? JJ] 1 «’ (Ibrahim 1933: 38, in Jansonius
& Hills 1976: 2362). It is clearly a rather insecure ‘portmanteau’ genus. Potonié &
Kremp (1954), quoted in Jansonius & Hills (1976), provided a more precise
diagnosis intended to accommodate a group of fungal spores.

The species described below conforms in its general features to Reticulata-
sporites (sensu lato), so that in the absence of a thorough revision of the genus and
its related forms it has been attributed to this genus, despite marked differences
from the type species of the genus.

Reticulatasporites grandis sp. nov.
Fig. 6E—-F

Etymology

The name of this species reflects the large size of the grain.

Description

The spores are atreme and spherical to subspherical—ellipsoidal. The whole
surface of the spore is covered by a large-meshed, quite regularly sized reticulum,
the muri formed apparently by steep folding of the outer skin. The muri
themselves are never straight but ‘wriggle’ across the surface. The lumina are
polygonal, mostly pentagonal, in shape. A pinnacle may be formed at the
junction of the muri. The diameter of the grains is 50-60 w.

Remarks

Reticulatasporites grandis may be more similar to other species of Reticulata-
sporites than it is to the type species of the genus. There is a superficial
resemblance between R. grandis and Retiperiporites piacabucuensis Herngreen
(1975b: 110, pl. 2 (fig. 5)), known from the Upper Senonian of Brazil.

Affinity

The affinities of Reticulatasporites grandis are open to speculation. It is
unlike lycopodiaceous spores. It is more likely that it has an affinity with a
bryophyte family, such as the Cleveaceae, whose members may also produce
atreme spores with large reticulate features (Boros & Jarai-Komlddi 1975).

Distribution

Reticulatasporites grandis is common in the Arnot samples. There is only
negative information about its wider distribution. It has apparently not been
recorded in Australian late Cretaceous and Tertiary sediments. A single specimen

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

encountered in late Cretaceous sediments off the south-western Cape coast and
described as ‘Lycopodiumsporites facetus’ appears from the photomicrograph
(McLachlan & Pieterse 1978: 875, pl. 2 (fig. 1)) to be indistinguishable from the
present form.

Spores not assigned to genus
Forma A
Fig. 7C-D
Few specimens of this spore were seen and it was unclear whether the robust
verrucate structures occurred on both faces. It is probably verrucate on the distal
face alone, which would place it in the genus Distaverrusporites, known from the

late Cretaceous of Nigeria and Borneo (cf. Van Hoeken-Klinkenberg 1966: 43,
pl. 1 (fig. 6)). Equatorial diameter 50 w, height of verrucae 6 wp.

Forma B
Fig. 8G-—H
Few specimens of this spore were seen. Equatorial diameter 32-34 wp.
Reticulum only on distal face. Muri 2 wu high. The junction of muri is marked by a
truncated spinate process, which is an additional 3—4 w higher than the muri. This
spore can probably be assigned to the genus Retitriletes, which has lycopodiaceous
affinities.

Forma C
Fig. 7E-F
Equatorial diameter 50 w. The spore is trilete. Laesurae are straight and two-
thirds the radius of the spore. Distal face is hamulate. Proximal face is covered by
a perine, loosely attached to the exine and tightly folded to form a fine rugulate—
hamulate pattern. Exine 3 w. This spore could perhaps be assigned to Hamulati-
sporis Krutzsch, 1959 (a subgenus of Camarazonosporites), which is known from
late Cretaceous and Eocene deposits in Europe (Azema & Ters 1971: 272).

Forma D
Fig. 7A-B
This is a very large trilete microspore with a massive perinous layer, up to
20 mw thick on the distal face and the equatorial region. This layer is coarsely and
jaggedly verrucate or fossulate. The proximal face is subpsilate. The laesurae are
thin and straight, or with a slight curve, and almost reach the equator. They are
placed on the apex of raised triangular ridges running the length of the radial
areas. This form distinctly resembles the spores of the extant dwarf tree-fern
genus Lophosoria (family Lophosoriaceae) as illustrated in Murillo & Bless
(1974: 252-253). This neotropical family is sometimes included in the
Cyatheaceae.

PALYNOLOGY OF THE ARNOT PIPE

eg Ae or eee

404

Fig. 7. A-B. Forma D. C-D. Forma A. E-F. Forma C (Hamulatisporis).

St

32

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. A. Araucariacites australis. B. Araucariacites sp. C-F. Zonalapollenites sp. B
(Cingulatipollenites). G—H. Forma B.

PALYNOLOGY OF THE ARNOT PIPE 35

Pollen of Coniferae
Inaperturate pollen
Genus Araucariacites Cookson ex Couper, 1953
Araucariacites australis Cookson ex Couper, 1953
Fig. 8A

Granulonapites (Araucariacites) australis Cookson, 1947: 130, pl. 13 (figs 1-4).
Araucariacites australis Cookson, 1947. Balme, 1957: 31, pl. 7 (figs 81-82). Kemp & Harris, 1977:
25, pl. 4 (figs 15-16).

Description

The pollen grains are large, always flattened and crumpled; inaperturate and
with a thin, finely granulate exine. Diameter 50-60 pw, exine 0,5 pw.

Affinity

See discussion in Cookson (1947: 130) and Kemp & Harris (1977: 25). The
affinity of A. australis is with the Araucariaceae and probably the genus
Araucaria.

Distribution

Araucariacites australis is common at Arnot, reaching relative abundances of
3 per cent. This species has a world-wide distribution in the Mesozoic, but is
mainly restricted to the Southern Hemisphere in the Tertiary. So far as is known
it is not present in the Neogene of the African sub-continent (Thiergart er al.
1963; Coetzee 1981). It is common in lower Cretaceous sediments off the east
coast of southern Africa (Scott 1976; McLachlan & Pieterse 1978), but less
common in the interior (Scholtz & Deacon 1982) and south-western Cape in
Cretaceous sediments. Together with other inaperturate forms, it dominates early
Palaeocene or late Cretaceous assemblages from the Gamoep area (unpublished
data).

Araucariacites sp.
Fig. 8B
Compare

Inaperturopollenites limbatus Balme, 1957: 31, pl. 7 (figs 83-84).
Balmeiopsis limbatus (Balme) Archangelsky, 1977: 122-126, pl. 1.
Araucariapollenites laffittei Reyre, 1973: 157, pl. 35 (figs 3-4).

Description

Outline oval or subcircular, exine 2-3 yw thick and granulate, diameter
57-74 w.

Discussion

The exine is much thicker than that of Avraucariactes australis but the
granulate sculpturing is very similar. Few specimens were seen in the samples
from Arnot, and although preservation of all specimens was excellent, doubt

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

about its morphology remains. Possibly what is here described as a thick exine, is
rather an equatorial crassitude or a particular concentric folding pattern of larger
specimens of the thinner-exined A. australis. Inaperturopollenites limbatus
Balme, 1957, is large, robust, granulate and inaperturate. Archangelsky (1977)
has recently instituted the genus Balmeiopsis for larger spherical, granulate grains
with an equatorial crassitude and an irregular aperture or thinning of the exine at
one pole. Reyre (1973) erected Araucariapollenites on the rather tenuous basis of
SEM-observed ultrasculptural exine features. Since few specimens were observed
at Banke and an irregular aperture was not noted, a definitive description of the
grain is left for later research. Palynomorph assemblages from other crater-lake
deposits in the same area, but somewhat earlier in time, consistently contain
A. australis and a larger and more robust morphotype.

Affinity

Araucariacites sp. is probably an extinct species of Araucaria. Balmeiopsis is
found in association with twigs and cones associated with Brachyphyllum-type
foliage, a leaf genus with affinity to the Araucariaceae (Archangelsky 1977).
Reyre (1973) compares Araucariapollenites to the pollen of Araucaria araucana
(cf. A. araucana Heusser, 1971: 12, pl. 8 (fig. 55)).

Distribution

Araucariacites sp. occurs in the Australian lower Cretaceous where it is
always rare (Balme 1957: 31; Burger 1973: 100). Balmeiopsis is recorded from
lower Cretaceous sediments of South America and Canada (Archangelsky 1977)
and Araucariapollenites from Mesozoic sediments of north Africa (Reyre 1973).

Monosaccate pollen
Genus Zonalapollenites Pflug, in Thomson & Pflug, 1953

See discussion in Dettmann (1963: 99, under Tsugaepollenites) and in
Jansonius & Hill (1976: 3093, 3265).

Zonalapollenites sp. A
Fig 9A—D
Description

The corpus is biconvex or boat-shaped in transverse section and the grain is
perisaccate in the equatorial—subequatorial region. The amb is roundly rectangu-
lar to broadly elliptical. Distal and proximal faces appear to have fused so as to
create a thick, laminated main body, which is granulate and rugose to
indeterminately sculptured. Initial folds in the exoexine on both the proximal face
(the face on to which the saccus overlaps) and the distal face mark the attachment
of the saccus. The saccus is formed from a relatively thick exoexine, which is
sometimes deeply folded in a radial direction and is distinctly less well developed
at the opposing poles of the longest axis of the amb. It would appear that the zone
of attachment of the saccus to the main body is subequatorial on the proximal and

PALYNOLOGY OF THE ARNOT PIPE 35

Fig. 9. A-D. Zonalapollenites sp. A. E. Lygistepollenites sp. F-1. Podocarpidites sp.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

perhaps also the distal face, resulting in a characteristic subequatorial dark ring
when the grain is viewed from the polar position. This effect is caused by the
density of the sporopollenin in the equatorial to subequatorial zone. Total
diameter 33-49 w, saccus width 9-17 w, diameter of the main body 28-38 yp,
thickness of main body 3—4 pw, height of the whole grain 22 wp.

Discussion

This species is similar to those described as Zonalapollenites segmentatus
(Balme 1957: 33, pl. 9 (figs 93-94)) and Tsugaepollenites segmentatus (Dettmann
1963: 101, pl. 24 (figs 6f, 11-16)) but differs clearly from the description of the
latter in that the present species does not have polar vesiculae. The present
species also differs in the characteristic wide, dark subequatorial (taking the grain
as a whole) ring. It is very difficult to photograph in transverse section. It must be
pointed out that the original generic diagnosis specifies that the “saccus’ is in fact a
velum formed by a fibrous baculate extension of the exine. Dettman’s (1963)
diagnosis of the genus mentions only an equatorial saccus and both Zonala-
pollenites sp. A and Zonalapollenites sp. B, described below, fit this more recent
diagnosis.

Affinity

Dettmann (1963: 100) quotes other authors in support of a coniferous
affinity — possibly to the genus Tsuga. Muller (1968) quotes Gamerro (1965) who
suggests a podocarpaceous affinity for Zonalapollenites.

Distribution

The genus is present world-wide in sediments of Jurassic to Palaeogene age.
It is rare in sediments of Cretaceous age off the southern African coast.
Zonalapollenites sp. A is rare at Arnot.

Zonalapollenites sp. B
Compare
Cingulatipollenites aegyptiaca Saad & Ghazaly, 1976: 449, pl. 13 (figs 3-6).

Description

The structure is complex. The grain is biconvex to flat. The amb is
subcircular to broadly elliptical. Monosaccate equatorially with the limbatus
marked by a distinct, sharp, irregular line. The saccus is of medium width and it is
not prominently folded in a radial or any other direction, but is rather
characterized by superficial pliae. The saccus is robust and may terminate
equatorially in a crassitude. The exine of the main body is relatively thin and
granulate proximally and distally. The outline of the grain is irregular and the
saccus may overfold the proximal surface, resulting in a rough frill-like line
internal to the limbatus. On occasional grains a trilete fold may mark laesurae.
The large size range makes it likely that more than one species is present.
Diameter 36-70 yw, width of saccus 6-12 wp.

PALYNOLOGY OF THE ARNOT PIPE a7

Discussion

The grains are different from those of Zonalapollenites sp. A in having a
relatively much smaller saccus, which is not folded in a radial direction. The
saccus in the present species also does not so regularly or prominently overfold
the proximal surface. The exine in the present species is granulate, while in
Zonalapollenites sp. A it is granulate—rugose. Although the outline of the grain is
irregular it does not show the undulate outline of Zonalapollenites dampieri, nor
has the overfolding of the saccus of the proximal face been mentioned as a
characteristic of Z. dampieri (Balme 1957: 32, pl. 8 (figs 88-90); Dettmann 1963:
100, pl. 24 (figs 1-5)). In other respects the present species is similar to
Z. dampieri.

Saad & Ghazaly (1976) have described Cingulatipollenites aegyptiaca from the
Nubia Sandstones of North Africa, which is almost certainly conspecific with
certain of the forms encountered in this study. However, some of the present
specimens do appear clearly saccate, rather than cingulate, so that the genus
Zonalapollenites is preferred in the present context. As indicated the size range and
morphological variability make it likely that more than one species is present, and
perhaps cingulate as well as saccate forms. Further study of these forms is required.

Affinity
Zonalapollenites sp. B is presumably coniferous, but there is no further
information on its specific affinities.

Distribution

Zonalapollenites sp. B is common at Banke. The genus is known world-wide
from the Jurassic to the Palaeogene. Cingulatipollenites is present in Jurassic to
Upper Cretaceous assemblages from the Nubia Sandstones, North Africa (Saad
& Ghazaly 1976).

Disaccate pollen
Genus Lygistepollenites Stover & Evans, 1973
Lygistepollenites sp.
rice IE
Compare

Dacrydium cupressinum Couper, 1953, pl. 4 (fig. 35).

Description

The grain is disaccate; the corpus is circular and covered proximally and
laterally with a wide layer of steep folds forming a rugulate pattern. The sacci are
small and half pendent on the distal face. The sacci seem to be formed merely by
larger folds of the outer skin. They are folded in a radial direction and there is no
distinct zone of attachment. The distal face bears less robust sculptural elements
and the sulcus, wide in the centre and narrowing towards the equatorial margins,
is clear.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Only one specimen in a good state of preservation was observed. The grain
lacks prominent proximal protuberances between the body and the proximal
roots of the sacci and thus does not belong to the genus Phyllocladidites Cookson
ex Couper, 1953, despite its small sacci. The grain is similar to grains of the extant
species Dacrydium cupressinum (Pocknall 1981: 70, figs 2a—e), although its sacci
may not be as well developed.

Affinity
The affinity of the genus Lygistepollenites is with the genus Dacrydium
(Section B) of the Podocarpaceae.

Distribution

Only one grain of Lygistepollenites sp. was observed at Arnot. The genus
Lygistipollenites is known from the Oligocene to the present in New Zealand. It
has not been recorded in late Cretaceous sediments from the interior of the
African subcontinent (Scholtz & Deacon 1982) nor off the Cape (McLachlan &
Pieterse 1978), but it is probably present in the Neogene (?) in the southern
(Thiergart et al. 1963) and south-western Cape (J. A. Coetzee, Institute for
Environmental Sciences, University of the Orange Free State, pers. comm.). It is
not recorded from late Neogene sediments from Burundi (Sah 1967).

Genus Podocarpidites Cookson ex Couper, 1953
Podocarpidites sp.
Fig. 9F-I
Compare

Disaccites grandis Cookson, 1953: 47, pl. 2 (fig. 41).

Pityosporites grandis (Cookson) Balme, 1957: 36, pl. 10 (figs 110-111).

Alisporites grandis (Cookson) Dettman, 1963: 102, pl. 25 (figs 1-5). Haskell,
1968: 217, pl. 1 (figs 1-2).

Description

The corpus is circular in polar view and the exine is thick. The outline of the
corpus is sometimes difficult to see. The sacci are large, semicircular and slightly
wider than the corpus. On the distal face their roots are clearly marked, spaced
wide apart and parallel, and outline a correspondingly broad tenuitas. The
reticulum of the sacci is coarse and often discontinuous towards the margins of the
sacci. The grain is sometimes apparently collapsed, in which case the tenuitas
appears narrower and tending towards fusiform while the shape of the grain
becomes oval. The preservation is usually poor. Length of the expanded grain
88-119 w, diameter of the corpus 60-63 pw, length of the sacci 68—75 ww, breadth
of the sacci 25-32 yw, distance between the zones of attachment on the distal face
25-33 p, exine of irregular height, 4-8 wp.

PALYNOLOGY OF THE ARNOT PIPE 39

Discussion

Haskell (1968: 217) provides a satisfactory description on the range of
variation in Alisporites grandis. The dimension of the grains that he measured are
very similar to those of Podocarpidites sp. He described two states for the
form—a diploxylonoid and a haploxylonoid state—which may correspond to the
‘expanded’ and ‘collapsed’ states described here. Despite regarding the present
form as conspecific with A. grandis (Cookson) Dettmann, 1963, as described by
Haskell (1968), and since the ‘expanded’ or diploxylonoid state (which shows
non-Alisporites-like characteristics) is most common at Banke, it was not
considered justified to place it in the genus Alisporites. It is beyond the scope of
the present paper to propose new combinations, and it is therefore merely noted
that the genus Alisporites Daugherty, 1941, has been redefined (Jansonius 1971;
Jansonius & Hill 1976: 68-69) and, in the opinion of the present author, the
diploxylonoid state of Podocarpidites sp. and of A. grandis as redescribed by
Haskell (1968) does not permit the inclusion of these species in the revised
diagnosis of the genus Alisporites. Jansonius (1971) also suggested a pterido-
spermous affinity for Alisporites, while the morphology of the present form
suggests a podocarpaceous affinity. The present form is probably comparable to
that described as Podocarpidites sp. by Sah (1967: 44, text-fig. 13, pl. 4 (fig. 11))
from the Neogene of Burundi.

Affinity

The affinity of Podocarpidites sp. is probably with the Podocarpaceae,
section Eupodocarpus (which includes the African species Podocarpus latifolius,
P. elongatus and P. henkelii) or section Stachycarpus (A. R. H. Martin 1959;
Pocknall 1981). On overall size alone the New Zealand members of what has
been regarded as the most primitive section of the family, Stachycarpus (Bucholtz
& Gray, 1948), compare most closely with the present form. The rarity and state
of preservation of the specimens precludes finer morphological comparisons.

Distribution

Alisporites grandis is known from the upper Jurassic and lower Cretaceous
strata in Australia and Canada (Haskell 1968) and from Cretaceous strata off
South America (Archangelsky & Gamerro 1967). It is common in the Australian
lower Cretaceous. It is not recorded in Cretaceous sediments of DSDP 361 off the
south-western Cape coast (McLachlan & Pieterse 1978), nor is it present in the
Knysna lignites (Thiergart et al. 1963). It is present in the Neogene of central
Africa (Sah 1967), where it is very rare. Podocarpidites sp. is rare at Arnot.

Podocarpidites kamiesbergensis sp. nov.
Fig. 1OA—D

Etymology
This species is named after the nearby Kamiesberg Mountains.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

eal
:

ce
”
|

Fig. 10. A-D. Podocarpidites kamiesbergensis sp. nov. E-J. Podocarpidites riembreekensis
sp. nov. K-L. Arecipites sp. B. M-—P. Arecipites sp. A.

PALYNOLOGY OF THE ARNOT PIPE 4]

Description

Small, disaccate pollen grains, the corpus trapeziform in shape, but
somewhat arched proximally. The semi-hemispheric sacci are attached laterally at
a low angle. The grain is often not fully expanded, in which case the sacci appear
more pendent. The exine is relatively thick proximally and laterally, and
vermiculate to foveolate. The distal tenuitas is relatively broad and parallel-sided
and the infrareticulation of the sacci is robust, clear and mostly perfect. In polar
view the sacci are as broad as the corpus. Corpus circular in polar view. Outline of
the sacci regular. Total length of expanded grain 36—41 uw; height of corpus in
lateral view 20 w, width of corpus 23-26 uw, length the same; breadth of sacci
10 pw, depth 16 pw, length of sacci 20-23 yw, distance between lines of attachment
of sacci on the distal face 8-10 yp.

Discussion

Podocarpidites kamiesbergensis is smaller in size than the smaller species of
Podocarpidites described in the available literature, such as P. congoensis Sah
(1967: 43, pl. 4 (figs 5-6, 9-19)). Podocarpidites kamiesbergensis may be the same
as P. knysnanus Thiergart, Frantz & Raukopf, 1963, described from the Knysna
lignites, but this form is inadequately characterized and illustrated (Thiergart er
al. 1963).

Affinity

The affinities of P. kamiesbergensis are with the Podocarpaceae, especially
the section Afrocarpus (including Podocarpus gracilior and P. falcatus) as defined
by A. R. H. Martin (1959). It would be difficult to distinguish between the pollen
of P. falcatus and Podocarpidites kamiesbergensis.

Distribution

Podocarpidites kamiesbergensis is common at Arnot, reaching a relative
abundance of 5 per cent. It is not present in the late Cretaceous sediments from
the interior (Scholtz & Deacon 1982) or off the south-western Cape (McLachlan
& Pieterse 1978), but is possibly present in the Neogene (?) Knysna lignites.
However, the description of P. knysnanus Thiergart, Frantz & Raukopf, 1963,
does not permit identification with the present species.

Podocarpidites riembreekensis sp. nov.
Fig. 10E—J
Etymology

This species is named after the nearby farm Riembreek, which is also the site
of several crater-lake deposits.

Description

Medium-sized disaccate pollen grains. The corpus has a rounded rhomboidal
shape in polar view and is trapeziform in side view. The sacci are laterally
attached at a low angle and are rigid, hemispherical and slightly wider than the

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

corpus in polar view. The infrareticulum of the sacci is greatly reduced so that
only isolated sections of muri remain. The proximal and lateral surfaces of the
corpus are covered by sharply defined, densely packed rugulate to verrucate or
vermiculate structures, and the exine thickens in the proximal area to form a
distinct cappa. Proximally each saccus is attached to the corpus by two
crassitudes, which are not always prominently protuberant. Such protuberances
are not so well developed around the remaining lateral and distal zones of
attachment, but are developed enough to form a thick rugose collar that
constitutes the roots of the sacci. The roots are marked, but least robust on the
distal face where the sacci are separated by a wide, parallel-sided tenuitas.

Total length of grain 40-64 w; width of corpus 23-33 mw, length of corpus
25-35 mw; length of sacci 25-39 w, depth of sacci approximately 14 w, breadth of
sacci approximately 25 w, exine of sacci approximately 1 wp.

_ Discussion

The grain has very distinctive morphology. The infrareticulum of the sacci is
so reduced that it may be entirely absent over large areas. The exine of the sacci is
relatively thick. In polar view the four proximal crassitudes are marked and
produce the characteristic rhomboidal shape of the corpus. The proximal and
lateral sculpturing is distinct and robust, and is truncated as the distal surface is
approached. The proximal protuberances are part of the robust collar that
attaches the sacci to the corpus, and differ from the more localized structures
described for Phyllocladidites. The much larger sacci also distinguish this species
from Phyllocladidites.

Affinity
The affinity of Podocarpidites riembreekensis is with the Podocarpaceae, but
perhaps not with any extant genus.

Distribution

Podocarpidites riembreekensis is common at Arnot. As far as is known this
form has not been recorded elsewhere.

Pollen of Angiospermae
Monoaperturate pollen
Monocolpate pollen
Genus Arecipites Wodehouse, 1933

‘The genus Arecipites Wodehouse, 1933 was emended [sic] by Anderson
(1960) to include only those reticulate monosulcate pollen grains which have,
among other characters, lumina less than 0,5 wu in diameter. Reticulate
monosulcate forms whose lumina width exceeds 0,5 w were referred by Anderson

to the genus Liliacidites Couper, 1953’—Chmura (1973: 104). This procedure has
been generally accepted and is the one followed here.

PALYNOLOGY OF THE ARNOT PIPE 43

Arecipites plectilimuratus Chmura, 1973
Fig. 11A—B
Arecipites plectilimuratus Chmura, 1973: 104, pl. 21 (figs 1-3).

Description

Monosulcate; elongate—ellipsoidal in polar view. The sulcus extends the
whole length of the grain and may even transgress the ends of the grain. The
margins of the sulcus are faint and irregular since the exine thins as the sulcus is
approached. The sulcus is open and, despite the irregularity, more or less
parallel-sided for its whole length. The exine is relatively thick and the reticulum
is distinct and uniform over the whole of the grain, except when the sulcus is
approached and the reticulum becomes indistinct. Muri are approximately the
same width as the lumina and mostly duplibaculate. Length of illustrated grain
40 pw, width 28 pw, exine 2 yp.

Discussion

For comparison with other species of Arecipites see Chmura (1973: 104). The
genus is a generalized morphological type and no specific identity of the plants
involved need be supposed.

Affinity

The affinity of Avicipites plectilimuratus is with a broad monocotyledonous
group including the Amaryllidaceae, Iridaceae and Liliaceae. The pollen of
southern African Monocotyledonae is not sufficiently well known to enable one
to suggest closer affinities with any taxa of the local flora.

Distribution

Arecipites plectilimuratus is rare at Arnot. It is also rare in the late
Cretaceous of California.

Arecipites sp. A
Fig. 1OM-—P

Compare

a

Arecipites reticulatus (van der Hammen) Anderson, 1960: 18.

Description

Small, monosulcate pollen, elongate—oval with rounded ends. The sulcus is
long, has no margo and reaches the ends of the grain. The exine is relatively thick
and clearly differentiated into a nexine and a tectate sexine. The surface of the
grain is uniformly covered by microscrobiculi. Length of the grain 23-25 yw, exine
1—2 p thick, scrobiculi 0,2 pw.

44

Fig. 11.

ANNALS OF THE SOUTH AFRICAN MUSEUM

A-B. Arecipites plectilimuratus. C-D. Liliacidites sp. B. E-F. Clavatipollen-
ites spp. (SEM). G-L. Clavatipollenites sp. A. M-N. Clavatipollenites sp. B.
O-Q. Clavatipollenites sp. C. R-T. Monocolpopollenites sp. B.

PALYNOLOGY OF THE ARNOT PIPE 45

Discussion

Arecipites sp. A is a small, robust, distinctive grain that is dissimilar to
A. plectilimuratus. It differs from A. reticulatus in its finer scrobiculi and thicker
exine.

Affinity
Arecipites sp. A has monocotyledonous affinities.

Distribution

Arecipites sp. A is rare at Arnot.

Arecipites sp. B
Fig. 1OK-L

Description

The grain is ellipsoidal with a thin and indistinct colpus, which runs the
length of the grain. The exine is relatively thick and the sexine and nexine layers
are clearly differentiated. The sexine is psilate to indeterminately sculptured and
appears finely columellate. Dimensions of illustrated specimen 38 X 26 w, exine
1-2 p.

Affinity
No information exists regarding the affinity of Arecipites sp. B.

Distribution

Arecipites sp. B is rare at Arnot.

Genus Liliacidites Couper, 1953
Liliacidites sp. A
Fig. 12E—H

Description

The grains are ellipsoidal to elongate—ellipsoidal in polar view. The sulcus in
expanded grains is wide with parallel sides, and stretches the length of the grain.
The meshes of the complex reticulum become smaller as the sulcus is approached
until a margo is formed by a narrow zone of unbroken exine. The exine may also
become thinner towards the colpus, but this was not definitely established.
Elsewhere the reticulum forms a complex, irregular, non-perfect pattern and is
simplicolumellate. The exine is relatively thick and two layers are clearly
distinguishable. Sexine and nexine are of similar height. Length of grain 29-36 p,
width of expanded grain in polar view 30 uw, sulcus width 3,5-5 w, exine
approximately 2 w, lumina 1-3 wp.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Liliacidites sp. A appears to be similar to those described as Liliacidites
intermedius Couper, 1953, but is distinguished from that species by the
characteristic reduction of the reticulum in the vicinity of the colpus, and in that
the reticulum of the present species is not reduced towards the ends of the grain.

Affinity

The affinity of Liliacidites sp. A is perhaps with the genus Chamaedorea of
the Palmae. Similar forms are also found in the Butomaceae and Liliaceae. The
extant genus Chamaedorea contains rather unusual small, reed-like palms found
in tropical deciduous thickets (Lozano-Garcia 1979).

Distribution

Liliacidites sp. A is common at Arnot.

Liliacidites sp. B
Fig. 11C-—D

Description

Monocolpate, irregular elongate—ellipsoidal. Narrow sulcus with distinct
margo stretches the full length of the grain. The tectum is imperfect.
Crassisexinous. The reticulum is irregular, imperfect and robust, so that the
sculpturing appears reticulate-rugulate. The columellae may be horizontally
elongated so that the sculpturing gives the appearance of broken stretches of
muri. Length of grain illustrated 36 w, width 22 pw, exine 3 w.

Discussion

Liliacidites sp. B is most unusual in the reduction in height of the columella
and a corresponding increase in the depth of the tectum.

Affinity
The affinity of Liliacidites sp. B is perhaps with the Liliaceae (cf. Lilium
longiflorum (Huang 1972: 266, pl. 173)).

Distribution

Liliacidites sp. B is very rare at Arnot.

Genus Clavatipollenites Couper, 1958

See discussions in Dettmann (1973: 11) and Kemp & Harris (1977: 55). The
genus is well represented at Banke with at least three forms being present, one of
which is common.

PALYNOLOGY OF THE ARNOT PIPE 47

Clavatipollenites sp. A
Fig. 11G—L
Compare
Clavatipollenites sp. Dettmann, 1973: 11, pl. 2 (figs 8-11).

Description

The grains are monocolpate and subspheroidal to slightly ellipsoidal—
spherical. The colpus is usually ulcerate with a broken and irregular margin in the
sexine. It is irregular in outline and usually more or less isodiametric. Occasional
grains have a clean, straight margin in the sexine (Fig. 111) and others appear
trichotomosulcate (Fig. 11J). Sexine and nexine are about the same height. The
reticulum is perfect and the muri simplicolumellate. The swollen heads of the
columellae and thus ‘lumpiness’ of the muri can be seen in the SEM
photomicrographs (Fig. 11E—F). Further SEM work may confirm whether the
‘lumpiness’ (Kemp & Harris 1977) or ‘beaded’ characteristics (Coetzee 1981) of
the exine can be of taxonomic significance. It would appear from the limited SEM
work done in the present study that in Clavatipollenites sp. A the ‘beadedness’ is
caused by suprategillar microconi mounted on the branches of the reticulum,
while in Clavatipollenites sp. C (described below) the ‘lumpiness’ of the muri may
be caused by the swollen heads of the columellae alone. Length of grain usually
between 22-27 uw, but occasional grains up to 31 mu, exine 1,5-3 w, lumina
approximately 1 mw, sulcus usually in the order of 5—8 w diameter, but may be
longer.

Affinity

See discussions in Dettmann (1973: 11), Kemp & Harris (1977: 56) and
Muller (1981: 9). Affinity of the genus Clavatipollenites is probably with the
Chloranthaceae (Doyle 1969).

Distribution

The distribution in time and space and a suggested pattern of extinction of
the Clavatipollenites—Ascarina complex is discussed in Muller (1981: 9-12). Their
suggested pattern of extinction needs to be revised since Clavatipollenites has
been recorded at Arnot and in the Neogene in the south-western Cape (Coetzee
1981). Observations from the Botswana region (Scholtz & Deacon 1982),
however, confirm that in late Cretaceous assemblages dominated by Ephedripites
(and Cretacaeisporites), Clavatipollenites is not present. Clavatipollenites sp. A is
common at Arnot.

Clavatipollenites sp. B
Fig. 1LM-N
Description

Monocolpate, subspherical to ellipsoidal—spherical. Sulcus relatively smaller
than in Clavatipollenites sp. A and more circumscribed. Exine thick and

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

crassinexinous. Recticulum relatively fine. Length 24 uw, width 19 w, nexine
approximately 2 w, sexine 1 w, lumina 1 wp.

Discussion
Clavatipollenites sp. B is distinguished from Clavatipollenites sp. A by the
prominent dense nexine, relatively thin sexine, and fine reticulation.

Distribution
Clavatipollenites sp. B is rare at Arnot.

Clavatipollenites sp. C
Fig. 110-Q

~ Description

Monocolpate, ellipsoidal—spherical with a long closed colpus, which, as usual
for Clavatipollenites, has ragged broken edges in the sexine. The exine is
relatively thick with sexine and nexine about the same height. Markedly robust
columellae support a perfect reticulum. Columellae are only placed beneath wide
areas of the tectum formed at the junction of individual branches of the tectum.
Length of grain 32 uw, width 26 w, exine 3-4 yw, lumina 1-2 p.

Discussion

Clavatipollenites sp. C is clearly distinguishable from the former two species
by its greater size, long sulcus, and robust columellae and tectum.

Distribution

Clavatipollenites sp. C is rare at Arnot.

Genus Monocolpopollenites Pflug & Thomson,
in Thomson & Pflug, 1953

See discussion in Nichols et al. (1973) and Jansonius & Hill (1976: 1691).

Monocolpopollenites sp. A
Fig. 12A—D
Compare
Monocolpopollenites sp. Jardiné & Magloire, 1963: 211-212, pl. 8 (figs 31-32).

Description

Monocolpate, subcircular to ellipsoidal in polar view and ellipsoidal in
equatorial view. The colpus has a complex structure; it is parallel-sided and
reaches the ends of the grain. The lips of the colpus are folded inwards and

PALYNOLOGY OF THE ARNOT PIPE 49

Fig. 12. A-D. Monocolpopollenites sp. A. E-H. Liliacidites sp. A. I-J. Forma E
(monocolpate pollen). K-—M. Milfordia hypolaenoides. N. Milfordia sp.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

thickened, and on the inner margins of the lips a scabrate row of sculpturing is
formed, consisting perhaps of closely packed, small verrucae. This is a very
characteristic feature. The exine is thick, tectate and psilate, and a finely punctate
perinous membrane envelops the grain. This layer sometimes closely adheres to
the surface of the grain and at other times separates from the exine. Few
specimens were observed, but from the size range it is possible that two species
may be present. Length 26-39 w, width 18-34 uw, exine 2,5—3 w, width of colpus
measured from the outer margins of the interior sculptured lips 5-8 w.

Affinity
The affinity of Monocolpopollenites sp. A is not known.

Distribution

Monocolpopollenites sp. A is rare at Arnot. Monocolpopollenites sp. Jardiné
& Magloire, 1963, is known from Turonian to Maastrichtian time ranges from
Senegal and the Ivory Coast.

Monocolpopollenites sp. B
Fig. 11R-T

Description

Grains monocolpate, rarely trichotomosulcate; ellipsoidal to subcircular.
The colpus stretches the length of the grain, is parallel-sided and relatively wide.
The lips are raised and decorated with short folds or verrucae. The ends of the
colpus are abruptly truncated and edged with verrucae or bits of exine. The exine
is relatively thin and is sparsely and irregularly decorated with isolated spinosa.
Length of grain 30-34 uw, width 20-23 uw, width of colpus approximately 4 uw,
exine approximately 1 wp.

Discussion

Monocolpopollenites sp. B is distinct from Monocolpopollenites sp. A in
having a much thinner exine and lacking a perinous layer. The latter may be a
weak criterion considering that only a few grains were observed. The sparse, but
always present, isolated spinosa of Monocolpopollenites sp. B are also distinctive.

Affinity

The affinity of Monocolpopollenites sp. B is perhaps with the Palmae,
although as far as could be ascertained no extant Palmae display similar
sculpturing or end-aperture morphology (Sowumni 1972; Kedves 1980).

Distribution

Monocolpopollenites sp. B is rare at Arnot.

PALYNOLOGY OF THE ARNOT PIPE Sl

Monoporate pollen
Genus Milfordia Erdtman, 1960
Milfordia hypolaenoides Erdtman, 1960
Fig. 12K—M
Milfordia hypolaenoides Erdtman, 1960: 46, pl. 1 (fig. a). Martin, 1973: 37, figs 163-165.

Description

The pollen is monoaperturate and spherical to subspherical. The pore is
relatively large, circular or elliptical, and ulcerate, with the margins ragged and
often with loose pieces of exine in the mouth. The pollen has a slightly irregular,
undulating outline and the exine is relatively thick, scrobiculate and possibly
finely fossulate. Diameter of grain 25-32 « on SEM photomicrographs, 31-43 yu
on the light microscope; aperture diameter 11-20 w, occasionally smaller and
clotted with bits of exine; exine 1-2 wp.

Discussion

Although Elsik (1968: 313) and Partridge (in Stover & Partridge 1973: 262)
proposed generic diagnoses for types of restionaceous pollens that would include
both the smaller, porate or graminoid-type aperture and the larger centro-
lepidoid-type aperture (Chanda 1966), there is little value in using such a broad
concept on a specific level, as Partridge has done in Milfordia homeopunctata
Partridge, in Stover & Partridge, 1973. Since Chanda’s work (1966) it has been
clear that a distinction between the two types is significant in terms of the
phylogeny and plant geography (see Johnson & Briggs 1981: 458).

The normal aperture in the Arnot specimens of Milfordia hypolaenoides is
relatively large and ulcerate and clearly of the centrolepidoid type (Chanda 1966).
The aperture is similar to the ‘Hypolaena’-type illustrated in Couper (1960: 62,
pl. 9 (figs 26—27)) and H. A. Martin (1978: 191, pl. 7 (fig. X)) although,
unfortunately, as Muller (1981: 105) points out, only one species of the genus
Hypolaena has such a pore.

Erdtman (1938) and Chanda (1966) have suggested, on the basis of the
pollen morphology of extant species, an evolutionary sequence beginning with
the ‘primitive’ Centrolepidaceae, through those Restionaceae with centrolepidoid
apertures and the Restionaceae with graminoid apertures, to the ‘advanced’
Flagellariaceae and Poaceae (Chanda 1966: 396). Following the work of Hochuli
(1979), Muller (1981: 105) has noted the importance of the ‘relatively small
rounded—porate aperture with a more or less irregular margin and an indistinct
annulus which is transitional between the centrolepidoid and graminoid aperture
type’ in the early fossil record of the Restionaceae. This type of aperture has been
named the Restio subverticillatus-type by Muller (1981) and corresponds to the
‘Restio’-type of H. A. Martin (1978: 191, pl. 7 (fig. W)). The latter is once again a

Sy ANNALS OF THE SOUTH AFRICAN MUSEUM

rather unfortunate choice of name by Martin as the R. subverticillatus-type does
not include many members of the genus Restio (Chanda 1966).

It is clear that the evolutionary sequence proposed by Erdtman (1938) and
Chanda (1966) is not supported by the fossil record. All three apertural states of
the Restionaceae (graminoid, centrolepidoid, and transitional) are at present
known from late Cretaceous or Palaeocene sediments (Hochuli 1979; Salard-
Cheboldaeff 1979; Muller 1981; present study). It seems, however, that none of
the early forms show the extreme development of either centrolepidoid or
graminoid apertures that can be found in some extant species. In particular the
sharply protruding graminoid apertural state is as yet known only from possibly
early Neogene sediments (Thiergart et al. 1963).

The present evidence, confirming the early existence of the centrolepidoid
apertural state, indicates that the phylogenetic relationships inferred by Johnson
. & Briggs (1981) for the Restionaceae may need revision.

Affinity

Chanda (1966) and Ladd (1977) discuss the morphology of the pollen of the
Centrolepidaceae, Restionaceae and Flagellariaceae. Most modern Australian
species of Restionaceae have the centrolepidoid (Chanda 1966) or ‘Hypolaena’-
type apertures (Couper 1960; H. A. Martin 1978), while almost all southern
African species have the graminoid type (Chanda 1966). The present form,
Milfordia hypolaenoides, is therefore morphologically similar to extant Australian
Restionaceae rather than to the majority of southern African species. However,
the pollen of one southern African genus, Thamnochortus (Chanda 1966;
H. P. Linder, Bolus Herbarium, University of Cape Town, pers. comm.), is
comparable to the present form. Thamnochortus is unusual in a number of other
respects amongst the southern African Restionaceae, and the indications from
taxonomic studies are that it has had a long, isolated evolutionary history in the
subcontinent (H. P. Linder, pers. comm.). Milfordia hypolaenoides is also similar
to the pollen of Centrolepsis and Gaimardia of the Centrolepidaceae, but because
of the modern distribution of this family, an affinity between it and
M. hypolaenoides is considered unlikely.

Distribution

Milfordia hypolaenoides Martin, 1973, has been recorded from the late
Cretaceous of North America (Jarzen 1978) and in the Lower Palaeocene of
Europe where it continues into Miocene time ranges (Muller 1981). In Australia
it may be present from the lower Eocene (Stover & Partridge 1973) and is
common in younger sediments. Restionaceous forms were not recorded in late
Cretaceous sediments off the south-western Cape coast (McLachlan & Pieterse
1978). However, typically southern African Restio-type graminoid forms only
were recorded, and in great abundance, from the Neogene (?) Knysna lignites
(Thiergart et al. 1963). Milfordia hypolaenoides is common at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 53

Milfordia sp.
Fig. 12N
Compare

Restioniidites homeopunctatus Hekel, 1972: 15, pl. 6 (fig. 30).
Restioniidites pascuali Archangelsky, 1973: 385, pl. 9 (figs 4-8).

Description

The grain is large, ellipsoidal and monoporate. The pore is relatively small
and circular and the margin neatly defined. The exine is of medium thickness,
finely scrobiculate, and the outline of the grain is smooth. The dimensions of the
single illustrated grain are 53 x 39 wy, the pore is 8 uw wide, the exine 2 uw thick.

Discussion

Milfordia sp. is distinguished from M. hypolaenoides by its aperture
morphology, smoother exine and ellipsoidal shape. The present form is similar to
the ‘Restio subverticillatus’-type discussed in Muller (1981: 105). Since only one
specimen was observed it was not assigned to either Restioniidites homeopuncta-
tus or R. pascuali, with which it is compared.

Affinity

The affinity of Milfordia sp. is thought to be with some southern African
Restionaceae, especially ‘Restio subverticillatus’ (see discussion on M. hypo-
laenoides).

Distribution

Pollen of the ‘Restio subverticillatus’-type is known from the Maastrichtian of
north Africa (Jardiné & Magloire 1963) and the Palaeocene of south and north
America and Europe (Muller 1981). In Australia it is known from lower Eocene
to Miocene sediments (Hekel 1972; Stover & Partridge 1973). Only one specimen
was observed at Banke.

Dicolpate pollen
Genus Dicolpopollis Pflanzl, 1956
Dicolpopollis sp.
Fig. 13A—D

Description

The grain is dicolpate (disulcate?) and ellipsoidal. The colpi are three-
quarters the length of the grain and have distinct margo. The colpi are usually on
opposite sides of the grain (Fig 13C—D) but are occasionally on the same face
(Fig. 13A—B). Crassisexinous. The reticulum is fine and regular and the lumina
are circular. Length 37-43 w, width 22-30 w, lumina approximately 0,5 yp.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

oN:

Fig. 13. A-D. Dicolpopollis sp. E-I. Triorites operculatus sp. nov. J—M. Triorites sphericus
sp. nov. N. Triorites harrissii. O-P. Proteacidites sp. A. Q-R. Proteacidites sp. B.

PALYNOLOGY OF THE ARNOT PIPE 35)

Affinity

Disulcate pollen is found in extant members of the Amaryllidoideae of the
Amaryllidaceae, in some genera of the Iridaceae, in the Tofieldieae of the
Liliaceae, and in the Palmae (Chmura 1973). However, the disulcate pollen of
Monimiaceae most closely resembles Dicolpopollis sp.

Distribution

Dicolpopollis sp. is more common towards the base of the Arnot sequence.

Monocolpate pollen not assigned to genus
Forma E
Fig. 12I-J

Description

The grain is monocolpate with a relatively thin exine and all the specimens
seen had an irregular amb but were not folded. The colpus has no margo and in all
specimens was on the edge of the grain. Reticulate; the reticulum is quite coarse
and duplicolumellate. The muri and lumina are of about the same width and the
columellae usually encircle a lumina. Dimensions of illustrated specimen
5 < 26 IL:

Affinity
Forma E grains resemble the pollen of the extant palm species Areca
warburgiana (Sowumni 1972, pl. 1).

Distribution

Forma E grains are rare at Arnot.

Triaperturate pollen
Triporate pollen
Genus Triorites Cookson ex Couper, 1953
See discussion in Muller (1968: 14).

Triorites operculatus sp. nov.
Fig. 13E-I
Compare
Triorites festatus Muller, 1968: 15, pl. 3 (fig. 10).

Etymology

The specific name refers to the presence of an operculum.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Triporate, spherical grains with an exine of medium width, which appears
subpsilate to granulate under the light microscope. Under the SEM it can be seen
that the exine is, in fact, psilate with minute, relatively closely spaced tuberculata
or spinules, giving the granular appearance. The pores are circular and are
surrounded by a low annulus of intermediate width, which is formed by the sexine
being slightly raised away from the nexine, i.e. forming a simple low vestibulum.
The vestibulum is so small and often inconspicuous that the grain can hardly be
described as vestibulate. The sexine may also thicken slightly in the region of the
pore. The pore has an operculum bearing the same minute tuberculata and
attached to a psilate underlying layer. The operculum is sometimes absent. The
exine is relatively thin and the grain is usually folded, this folding following no
particular pattern. Two layers are distinguishable in the exine. Diameter
. 27-30 mw, exine approximately 1 w, diameter of pore approximately 2 uw, width of
annulus 1 p.

Discussion

Triorites operculatus is different from T. festatus Muller, 1968, in having an
operculum. The pores and annulus of 7. operculatus are also smaller.

Affinity

Muller (1968: 15-16) has warned that in the case of forms that show a
generalized, primitive type of pore (as in 7. operculatus), affinity with an extant
family will be almost impossible to determine with any certainty. Broad affinity is
possibly with certain families of the Hamamelidales (Takhatajan 1969), including
the Ulmaceae, Carpinaceae, Corylaceae, Casuarinaceae and Myricaceae. Oper-
culate forms are not uncommon in these families; all the Carpinaceae (some of
which are triporate), some species of the genus Celtis of the Ulmaceae (e.g. Celtis
iguanea—Erdtman 1952), and some of the Myricaceae are operculate.

Distribution

Triorites operculatus is the dominant form at Arnot, constituting 54 per cent
in one sample, and sometimes occurring in clusters. Triorites festatus is present
from the late Cretaceous in Borneo, but is more frequent in the late Palaeocene—
Eocene and follows the same pattern in tropical west Africa (Salard-Cheboldaeff
1981). The genus Celtis is known from leaf impressions of Maastrichtian age from
the Cameroons (Salard-Cheboldaeff 1981).

Triorites sphericus sp. nov.
Fig. 13J-M
Etymology

The specific name refers to the robust spherical shape of the grain and the
fact that the grain is seldom deformed.

PALYNOLOGY OF THE ARNOT PIPE Sy

Description

Triporate grain, spherical and robust. The exine is relatively thick and
crassisexinous. The pores are circular to equatorially elongated and are
surrounded by a clear annulus. The point at which the annulus begins is marked
by a sharp line of the exine surface. The annulus is formed by the sexine
separating and rising abruptly away from the nexine. The exine is subpsilate to
faintly rugose and undulate. In some grains it appears that the exine may thicken
slightly towards the mesocolpia in the equatorial region. Equatorial diameter
20-24 w, diameter of pore opening approximately 2 w, diameter of pore and
surrounding annulus 6—7 pw, exine 2-4 w.

Discussion

Triorites sphericus differs from T. operculatus in its smaller size, relatively
thicker crassisexinous exine, lack of operculum and in sculpturing details of the
exine. Unlike T. operculatus the grain of T. sphericus is seldom folded in any way.

Affinity
See note on affinity of Triorites operculatus. Triorites sphericus 1s perhaps

morphologically closer to species of the Betulaceae or Corylaceae than to those of
the Ulmaceae, Carpinaceae or Casuarinaceae.

Distribution

Triorites sphericus is common at Arnot.

Triorites harrissit Couper, 1960
Fig. 13N
Driorites harrissii Couper, 1960: 67, pl. 12 (fig. 2). Hekel, 1972: 17, pl. 5 (fig. 7).

Description

Triporate, angulaperturate with triangular amb. Exine is relatively thick with
nexine and sexine distinct. Subpsilate, pores narrow. Equatorial diameter of
illustrated specimen 31 yw, exine 2 w, pore diameter 2 w.

Discussion

Triorites harrissii differs in shape from the two previous species. The Arnot
specimens are very similar to the illustrated specimens of Triorites harrissii of
Couper (1960) and Hekel (1972).

Affinity
The affinity of TJ. harrissii is with the Casuarinaceae or Myricaceae—more
likely the former.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Triorites harrissii has a range from the Palaeocene to the present in the
Australia-New Zealand area. Only three specimens were observed at Arnot.

Genus Triporopollenites Pflug & Thomson,
in Thomson & Pflug, 1953

There is some confusion as to what generic designation should be used for
forms with the general morphology of the species described below. The problem
arises in that four genera have been used to describe broadly similar morpho-
types. These are Proteacidites Cookson ex Couper, 1953, Triporopollenites Pflug
& Thomson, in Thomson & Pflug, 1953, Echitriporites van Hoeken-Klinkenberg,
1966, and Propylipollis Martin & Harris, 1974. A. R. H. Martin & Harris (1974)
have also discussed the problems raised by the burgeoning of the genus
_ Proteacidites and proposed two additional genera—the three being distinguished
mainly in apertural morphology.

Examples of forms that are broadly related morphologically and have been
classified under the above four genera, are the following: Proteacidites tuberculi-
formis Harris, 1965 (p. 92, pl. 29 (figs 5—7)); P. longispinosus Jardiné & Magloire,
1963 (p. 218, pl. 7 (figs 15-17)); Echitriporites trianguliformis van Hoeken-
Klinkenberg, 1966 (p. 21, pl. 42 (fig. 7)); and Triporopollenites ambiguus Stover,
in Stover & Partridge, 1973 (p. 269, pl. 21 (fig. 7)). Proteacidites tuberculiformis
Harris, 1965, was later transferred by A. R. H. Martin & Harris (1974) to
Propylipollis although, according to Harris’s original description, it lacks the
diagnostic post-atrium. Martin & Harris also excluded Proteacidites longispinosus
from Proteacidites Cookson ex Couper, 1953 (sensu Martin & Harris), without
proposing an alternative genus. To add to the confusion Boltenhagen (1978)
instituted a new species Proteacidites sigalli Boltenhagen, 1978, which he
compared to Echitriporites trianguliformis, and suggested, on very poor grounds,
that P. sigalli could be compared with pollen of the extant proteaceous genus
Spatalla.

No new genus is proposed here but it is suggested that a genus might be
considered that would include medium to large triporate forms with simple pore
structures, whose sculpturing is of scattered apicula, spinosa or micro-echina, but
which are not echinate, i.e. forms comparable, for example, to the pollen grains
of the extant proteaceous genera Telopea and Embothrium (Erdtman 1952: 356).

Triporopollenites namaquensis sp. nov.
Fig. 14A-E
Compare

Triporopollenites ambiguus Stover, in Stover & Partridge, 1973: 269, pl. 21

(fig. 7).
Proteacidites tuberculiformis Harris, 1965: 92, pl. 29 (figs 5-7).

PALYNOLOGY OF THE ARNOT PIPE

S04

Fig. 14. A-E. Triporopollenites namaquensis sp. nov. F-J. Propylipollis meyeri
sp. nov.

a9

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The species is named after the region Namaqualand, which in turn takes its
name from its indigenous inhabitants, the Nama.

Description

Triporate, angulaperturate pollen with a triangular amb and straight to
slightly convex sides. The grains are large and, relative to their size, thin-walled,
so that they are mostly irregular, flattened and folded. Sexine and nexine can be
distinguished and the exine is crassinexinous. The thin exine is finely punctate and
bears solitary spinules mounted on broader bases, sparsely but more or less
regularly distributed on its surface. The pollen is very variable in size and this
suggests the possible presence of more than one species. The exine thickens
slightly at the pore margins to form a distinct annular ring, which is, however,
difficult to observe in polar view. The pore is circular to equatorially elongated.
Equatorial diameter 45-90 w, pore diameter 6-10 uw, width of annulus approxi-
mately 2 uw, height of spinosa 2 pw.

Discussion

A few species of broadly similar morphology are known (see discussion on
genus). Triporopollenites namaquensis differs from Proteacidites tuberculiformis
in being spinulate (not verrucate), in the sparseness of its ornamentation, and in
having a distinct annulus—although, as stated, it is easy to miss this feature.
Except for size, the present species being considerably larger, 7. namaquensis is
very similar to T. ambiguus.

Affinity
Triporopollenites namaquensis perhaps has affinities with members of the
subfamily Grevilleoideae of the Proteaceae.

Distribution

The species is common at Arnot. Germeraad et al. (1968: 312) in discussing
the distribution of the similar morphotype, Echitriporites, stated: “The more
triangular grains with fewer and smaller spines are more common in the Upper
Cretaceous of northern South America, whereas the more rounded grains with
slightly more and larger spines are more common in the Eocene.’ The similar
Australian form 7. ambiguus is known from Palaeocene and Eocene sediments.

Genus Proteacidites Cookson ex Couper, 1953

The generic name is used here in the restricted sense as defined by A. R. H.
Martin & Harris (1974). See also discussion for Triporopollenites (p. 58).

PALYNOLOGY OF THE ARNOT PIPE 61

Proteacidites sp. A
Fig. 130-P

Description

Small, colpoidate pollen (as for example Beauprea elegans (Erdtman 1952:
343)) with triangular amb. Angulaperturate. The exine is intermediately thick,
and thins at the margins of the colpi. Sexine thinner than nexine. Reticulate.
Diameter of illustrated specimen 22 yp, exine 1,5 yp.

Proteacidites sp. B
Fig. 13Q-R

Description

Small, triporate pollen with triangular amb; a rectangle with rounded corners
in equatorial view. The pore is small and circular and the pore margins are simple.
The exine is relatively thick, crassinexinous and reticulate. Diameter of illustrated
specimen 20 w.

Affinity
Pollens similar to Proteacidites sp. B are found in the Proteaceae and the
genus Allophylus of the Sapindaceae.

Distribution

Proteacidites sp. B is rare at Arnot.

Genus Propylipollis Martin & Harris, 1974
Propylipollis meyeri sp. nov. 7
Fig. 14F-J

Etymology
This species is named for A. P. and Christine Meyer of the farm Banke.

Description

Pollen triporate, angulaperturate, amb triangular, sides straight to slightly
concave, apices roundly truncate. Exine relatively thick and crassinexinous.
Nexine thickens as the pore is approached but is truncated before the pore in the
sexine to form a post-atrium (sensu Kremp 1965, fig. 380); the state of
preservation affects the visibility of this feature. Short radiating costae pori are
present. Sexine reticulate. The reticulate pattern is irregular, angular and not
always perfect and the size of lumina decreases towards the pores, so that in the
vicinity of the pores a foveolate structure exists. Loose pieces of sexine may
sometimes be present at the pore entrance. The coarseness of the reticulum varies

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

considerably and two species may be present. The muri are mostly simplicolu-
mellate, but in coarser areas duplicolumellate sections occur. The outline of the
grain is smooth. Equatorial diameter 25—33 uw, exine 2 uw, pore 2—4 w, lumina
approximately 1 w.

Affinity

The pollen of a number of genera of the subfamilies Grevilleoideae and
Persoonioideae of the Proteaceae have similar pollen. The pollen of southern
African species of Persoonioideae are not sufficiently well known to confirm a
closer affinity, but affinities to the genera Lomatia and Leucospermum have been
suggested (Germeraad et al. 1968).

Distribution

The species is common at Arnot. A related morphological form Proteacidites
dehaani Germeraad, Hopping & Muller, 1968 is common in the uppermost
Cretaceous and lowest Palaeocene strata of tropical Africa (Germeraad et al.
1968). The form is not present in late Cretaceous sediments off the south-western
Cape coast (McLachlan & Pieterse 1978) A probably conspecific form occurs in
the Neogene (?) Knysna lignites.

Propylipollis sp.
Fig. 1SA-B

Description
Triporate, the pores slightly protuberant and gaping. Angulaperturate. The
amb is triangular and the shape oblate. The exine is thick and has a complex
structure. Nexine, sexine and further subdivisions of the exine are clearly
differentiated. The sexine is thick, atectate and irregularly foveolate. The grain is
robust. Diameter of single specimen 44 w, pore 9-11 pw, exine 3-4 mw.
Only one specimen was seen and the above description should be regarded as
provisional.

Affinity
The affinity of Propylipollis sp. is perhaps with the tribe Grevilleeae of the
Grevilleoideae (Proteaceae) or with the Onagraceae.

Distribution

One specimen of Propylipollis sp. was observed at Arnot.

Tricolpate pollen
Genus Tricolpites Cookson ex Couper, 1953
See discussion by Kemp & Harris (1977: 29).

Fig. 15.

PALYNOLOGY OF THE ARNOT PIPE 63

A-B. Propylipollis sp. C-—G. Tricolpites reticulatus. H-J. Spinitricolpites
Jennerclarkei sp. nov.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tricolpites reticulatus Cookson, 1947
Fig. 15C-G

Tricolpites reticulata Cookson, 1947: 134, pl. 15 (fig. 45).
Tricolpites waiparensis Couper, 1960: 66, pl. 11 (figs 13-15).
Tricolpites reticulatus Cookson, 1947. Kemp & Harris, 1977: 30, pl. 5 (figs 1-2).

(See discussion by Kemp & Harris (1977) and Muller (1981: 67)).

Description

Tricolpate, fossaperturate; colpi short, extending about half-way to the
poles. The grain is circular in equatorial view, the amb is lobate, and the whole
surface is finely reticulate. The SEM photomicrographs illustrate the range of
variation in morphology. In Figure 15F the amb 1s distinctly more lobate and
fossaperturate, and the reticulate sculpture tends towards a tectate and foveolate
state; despite a narrow size range, two species may be present. The margins of the
colpi are marked by a flat seam in the sexine and a definite extension of the nexine
beyond the sexine at that point and into the mouth of the aperture. This can be
observed under both the light microscope and SEM. The grain is robust;
equatorial diameter 19-26 w, polar diameter approximately 20 uw; exine clearly
two-layered and approximately 1 mw; lumina 0,2—0,5 w.

Affinity

Tricolpites reticulatus is thought to have affinities with the genus Gunnera.
According to the data provided by Jarzen (1980) the Banke forms are the smallest
fossil Gunnera forms yet recorded, and closest—when compared to the average
for extant pollen from certain geographical regions—to those of South America
(acetolysed grains measured).

Distribution

Tricolpites reticulatus is common at Arnot, reaching a relative abundance of
7 per cent. It is known worldwide from the middle Cretaceous to the present and
is often common. See Jarzen (1980) for a full discussion of the occurrence of
Gunnera pollen in the fossil record as well as notes on the present-day habitat
requirements and distribution of the genus. A single species Gunnera perpensa, a
semi-aquatic species, is widespread in southern Africa except in South West
Africa—Namiubia.

Tricolpites sp. A
Fig. 16A-B

Description

Tricolpate; the amb is circular and the shape prolate. The colpi are long,
thin, simple slits that stretch three-quarters of the polar axis of the grain. The
exine is relatively thick and nexine and sexine are clearly differentiated. The
sexine is columellate and tectate, and the columellae are intermediately robust.
Dimensions 30-27 jw X 25-22 mw, exine approximately 1,5 w.

PALYNOLOGY OFRAHE ARNOT PIPE

Se SNE: OOS mess BZ

Fig. 16. A-B. Tricolpites sp. A. C-E. Crototricolpites densus. F-—G. Tricolpites sp. B.
- H-I. Tricolpites sp. C. K-—M. Tricolpites sp. D.

65

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity
There is no information regarding the affinity of Tricolpites sp. A.

Distribution

This species was infrequent at Arnot.

Tricolpites sp. B
Fig. 16F-G

Description

Tricolpate; the amb is triangular with convexly curved sides and angulapertu-
rate. No specimen was observed in equatorial view, but it appears that its shape is
_ biconvex. The colpi are very short, narrow slits. The exine is of intermediate
height and the sculpturing is verrucate to dispersed rugulate, producing a
negatively reticulate pattern.

Distribution

Tricolpites sp. B is rare at Arnot.

Tricolpites sp. C
Fig. 16H-I

Description

Tricolpate; the amb is triangular with convexly curved sides and angulapertu-
rate. No specimen was observed in equatorial view, but it appears that the shape
is flat and very slightly biconvex. The colpi are very short, narrow slits. The exine
is of intermediate width and granulate, and apparently thins in the immediate
vicinity of the colpi to form an almost exineless rim of regular width around the
colpi. Diameter of illustrated specimen 35 w.

Discussion

In the amb, position of the apertures and short, slit-like colpi there is some
similarity between Tricolpites sp. B and Tricolpites sp. C. Only two grains of each
were seen so that their descriptions must be regarded as provisional.

Affinity

There is a tenuous similarity between Tricolpites spp. B and C and the pollen
of certain extant members of Protea such as P. mellifera or P. grandiflora
(Erdtman 1952: 350).

Distribution

Tricolpites sp. C is rare at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 67

Tricolpites sp. D
Fig. 16K-M

Description

Tricolpate, amb circular to subcircular, shape oblate to biconvex. The colpi
are of intermediate width at the equator, relatively long, and narrow to a point at
their extremities. The exine is thick, and sexine and nexine are of similar width.
The nexine forms a uniform dense layer, which broadens equatorially at the colpi
margins to form marked costae endocolpi. This crassitude is apparently only
present equatorially. The sexine is of regular width and curves over the nexine at
the mouth of the colpi. The sculpturing is complex and coarse, and consists of an
irregular granulate surface covered by micro-echinae. Equatorial diameter
43-45 pw, exine 2,5—4 wu, colpi width 2,5-4 wp.

Affinity

In form, size and the unusual surface sculpturing there is some resemblance
between Tricolpites sp. D and the genus Ferocactus of the Cactaceae; see
Ferocactus latispinus (Lozano-Garcia 1979: 310, pl. 6). However, present-day
distribution and the established fossil record of the Cactaceae make this suggested
affinity somewhat unlikely and forms similar to Tricolpites sp. D occur in a
number of other families. The pollen is not similar to that of Rhipsalis, the only
extant genus of Cactaceae that is possibly indigenous to Africa.

Distribution

Tricolpites sp. D is infrequent at Arnot.

Genus Crototricolpites Leidelmeyer, 1966
Crototrico.pites densus Salard-Cheboldaeff, 1978
Fig. 16C-E
Crototricolpites densus Salard-Cheboldaeff, 1978: 224, pl. 1 (figs 10-12).

Description

The poilen grain is tricolpate, and almost circular in polar view. The grain is
invariably flattened. The colpi are broad equatorially and have gaping, ragged
margins; verrucae are the sculptural element and are angular, mostly triangular,
in polar view. Their tops are pointed, with groups of five to six arranged in a
circular pattern, the unit of a ‘croton pattern’. In polar view the colpus extends
about one-half the diameter of the grain. The equatorial diameter is 34 uw, width
of colpi at equator approximately 10 w, width of verrucae 1-2 yw, height of
verrucae approximately 1 w.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

The Banke specimens are similar to C. densus Salard-Cheboldaeff, 1978 and
differ from C. annemariae Leidelmeyer, 1966, in the relatively smaller verrucae
and circular ‘croton pattern’ units.

Affinity

The affinity of C. densus is with the Klaineanthus-type and perhaps, because
of the doubtful feature of ragged colpi margins, with the Adenocline-subtype of
the Crotonoideae of the Euphorbiaceae (Punt 1962).

Distribution

Crototricolpites densus is common at Arnot. The genus is known from the
lower Eocene of Guyana and the Oligocene and lower Miocene of tropical
Africa. Similar forms have not been recorded from Australia.

Genus Spinitricolpites gen. nov.

Diagnosis

Medium to large-sized, more or less spherical, tricolpate spiniferous pollen
grains. The spines are medium-sized, with pointed or rounded tops and are
sparsely and irregularly arranged on the sexine.

Type species Spinitricolpites jennerclarkei sp. nov. by original designation.

Etymology

The name refers to the spiniferous and tricolpate nature of the pollen grains.

Discussion

Apart from Tricolpites latispinosus McIntyre, 1965 (p. 207, figs 13-15) no
similar forms have been encountered in the literature. The combination of large
size, spherical shape, tricolpate state and spinate sculpturing would seem to
justify the erection of a new genus. Spinitricolpites gen. nov. is here considered to
include Spinitricolpites jennerclarkei sp. nov. and Tricolpites latispinosus
McIntyre, 1965.

Spinitricolpites jennerclarkei sp. nov.
Fig. 1SH-J

Etymology

The species is named after Mr Hugh Jenner-Clarke, an exploration geologist,
who during many years work has located numerous kimberlite pipes in the
Gamoep area.

PALYNOLOGY OF THE ARNOT PIPE 69

Description

The pollen grains are medium-sized, spherical to prolate spheroidal,
tricolpate and spiniferous. The colpi are straight slits with no margo and are about
one-half to two-thirds the polar diameter of the grains. In polar view the grain is
circular but due to flattening the colpi are open at the equator. The margins of the
colpi are not strengthened in any way and may appear frayed in polar view. The
possibility that the grains are split rather than tricolpate can probably be excluded
because of the regular positioning of the colpi in polar view and because the colpi
can be seen in equatorial view. The spines are medium sized, mostly with pointed
tips, and are sparsely and irregularly arranged on the surface of the grain; they
sometimes occur in clumps or irregular rows. The exine is clearly differentiated
into a sexine and nexine and the sexine is tectate, finely columellate and punctate.
Equatorial diameter excluding spines 39-46 w, height of spines 3-5 pw, exine
1,5-2,5 w.

Discussion

The present form differs from S. latispinosus (McIntyre, 1965) in its longer,
more sparsely distributed spines.

Affinity

There is no information regarding the affinity of S. jennerclarkei. In the large
spherical shape and spinate sculpturing there is some resemblance to certain
members of the Valerianaceae or Verbenaceae (Huang 1972).

Distribution

Spinitricolpites jennerclarkei is common at Arnot; S. latispinosus (McIntyre,
1965) is known from the Miocene of New Zealand.

Tricolporate pollen
Genus Tricolporopollenites Pflug & Thomson, in Thomson & Pflug, 1953
Tricolporopollenites grandis sp. nov.
Fig. 17K-N

Etymology

The specific name refers to the large size of the grain.

Description

The morphology of the pollen is complex. It is large, tricolporate, striate, ora
lalongate or zonorate, highly prolate, ellipsoidal or with blunt ends. The colpi are
narrow and extend almost to the poles. The exine is thick and a robust, striate
sexine is clearly differentiated from the nexine. The nexine more than doubles its
width as the lalongate endoporus is approached to form a costae endoporus, and
the sexine thins in the small apocolpium. The striae are supported by wide

70

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. A-C. Tricolporopollenites brinkiae sp. nov. D-H. Tricolporopollenites arnotiensis
sp. nov. K-—M. Tricolporopollenites grandis sp. nov. O-S. Tricolporopollenites sp. A.

PALYNOLOGY OF THE ARNOT PIPE Wil

columella and sections of muri of the same width, arranged in an irregular
reticulate pattern that obscures the striate surface (which is so clearly visible in
the purely topographic SEM photomicrograph (Fig. 17N)). There seems to be a
centre line running length-wise down the middle of the mesocolpium, with a node
on the equator around which the shallow curving pattern of the striae is centred.
The colpi form a tangent to the arc of the striae with the position of the colporus
being the point of intersection of the colpi and arc. Also, the elements of the
striae are longest in the mesocolpium and apocolpium areas and are broken up
into shorter elements in the vicinity of the colporus. Length of polar axis 45-56 pw,
exine 2—5,5 w (including costae endopori), width of costae endopori 2,5-3,5 p,
sexine approximately 2 uw, width of lalongate ora or pore zone 2-3 w.

Discussion

This is an unusual grain, which is not really comparable to any known fossil.

Affinity

In size, shape, general aperture type and the complex morphology of the
sculpturing, the grain resembles the pollen of extant members of the genus
Bauhinia of the Caesalpinaceae (Senesse 1980: 394).

Distribution

Tricolporopollenites grandis is common at Arnot.

Tricolporopollenites arnotiensis sp. nov.
Fig. 17D-H

Etymology
This species is named after the Arnot Pipe.

Description

Tricolporate, amb tending towards lobate, planaperturate grain. Subprolate
in shape. The colpi are thin and about two-thirds the polar axis of the grain. The
ora are lalongate, wider than the colpi and relatively long. Sexine and nexine are
not always clearly differentiated, but the grain is crassisexinous. The sexine is
tectate and finely columellate and there may be fine suprategillar sculpturing.
Equatorial diameter 20—26 uw, polar diameter 20-24 w, ora width 1-2 w, exine
2 p, length of ora 6-7 w.

Affinity

There is some resemblance between 7. arnotiensis and the pollen of
Anthospermum (Rubiaceae). Tricolporopollenites arnotiensis also resembles
forms of the Euphorbiaceae with the Hippomane-configuration (Punt 1962), such
as Euphorbia hypericifolia and E. heterochroma illustrated in Bonnefille &
Riollet (1980, pls 47—48).

2: ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Tricolporopollenites arnotiensis is rare at Arnot. The extant genus Antho-
spermum is found in Africa and Madagascar.

Tricolporopollenites brinkiae sp. nov.
Fig. 17A—C

Etymology
The species is named after Brink Scholtz.

Description

The grain is tricolporate and robust. In overall shape the amb is lobate.
_ Fossaperturate. If, however, the sexine—which is very much thickened in the
equatorial mesocolpium—1is excluded, the amb of the remainder of the grain is
that of a very rounded triangle and the grain angulaperturate. In equatorial view
the grain is oblate. The sexine and nexine are clearly differentiated and the
prominent sexine is columellate, tectate and finely reticulate. The sexine thins
abruptly in the vicinity of the colpi and then bends upwards to edge the colpi with
distinct lips. The sexine is also thin in the apocolpium and the columellae are finer
here than elsewhere. Equatorial diameter 24 uw, width of colpi 2-3 w, nexine
approximately 0,5 uw, width of sexine in the mesocolpium 3 wp.

Discussion

Tricolporopollenites brinkiae has a very distinctive morphology and as far as
is known no fossil species closely resembles it.

Affinity

This species bears some resemblance to the pollen of the southern African
genus Nenax of the family Rubiaceae and, as with 7. arnotiensis, to certain
Euphorbiaceae (cf. Euphorbia heterochroma—Bonnefille & Riollet (1980,
pls 47—48)).

Distribution

Tricolporopollenites brinkiae is common at Arnot.

Tricolporopollenites coetzeeae sp. nov.
Fig. 18I-L

Etymology

This species is named after Prof. J. A. Coetzee, who has done pioneering
work on the Tertiary palynology of southern Africa.

PALYNOLOGY OF THE ARNOT PIPE

Fig. 18. A-B. Tricolporopollenites sp. C. C-F. Tricolporopollenites sp. D.
G-H. Tricolporopollenites sp. B. I—L. Tricolporopollenites coetzeeae sp. nov.
M-O. Forma H.

13

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Large, tricolporate and verrucate grain. The exine is relatively thin and
sexine and nexine layers are difficult to distinguish, except in the region of the
thin costae endopori where the nexine is prominent. The verrucae are spherical
and large relative to the exine. However, they are decorated with microconi that
give them an angular appearance. The colpi are narrow and almost reach the
poles of the grain. The ora are broad, lalongate and short and the costae endopori
are marked. The amb is subcircular to lobate. The grain is prolate but due to the
thin exine most specimens are deformed to some extent. Length 47—52 yu, width
30-34 w, exine 2-3 w including verrucae, width of ora 4-6 pw, length of ora
9-10 pw.

Affinity

In aperture morphology, size and general exine structure there is a similarity
_ between this species and the pollen of the genus Zimmermania (subfamily
Phyllanthoideae of the Euphorbiaceae) (Punt 1962: 29, pl. 4 (fig. 1)). However,
the prolate shape of T. coetzeeae, as well as the close packing of the verrucae and
their decoration with microconi, is a condition not approached by the pollen of
any of the seven extant species of Zimmermania. The fossil form is most similar to
the pollen of the species Z. acuminata, Z. ovata and Z. capillipes (Poole 1981).

Distribution

Tricolporopollenites coetzeeae is infrequent at Arnot. The genus Zimmer-
mania is endemic to East African montane areas, where it occurs mainly in mist
forests (Poole 1981).

Tricolporopollenites sp. A
Fig. 170-S

Description

The pollen is tricolporate, the shape prolate to pointed oval, fossaperturate
and striate, the amb is circular. The colpi are fairly deep and wide, and narrow
towards the poles. The ora are lalongate and pinched at the transection with the
colpi. There are marked costae pori. The striae are simplicolumellate. Equatorial
diameter of illustrated specimen 23 w, polar diameter 32 w.

Affinity
The affinity of Tricolporopollenites sp. A is probably with the genus Rhus of
the Anacardiaceae.

Distribution

The earliest records of pollen of the Rhus-type are from the Maastrichtian of
North America and upper Palaeocene of Europe. It has not been reported from

PALYNOLOGY OF THE ARNOT PIPE 1S

the early Tertiary of Australia and is not present in Maastrichtian sediments of
southern Africa (McLachlan & Pieterse 1978). It is present in the late Neogene of
central Africa (Sah 1967) and the Neogene (?) Knysna lignites. Members of the
Anacardiaceae are known from the Eocene of tropical Africa.

Tricolporopollenites sp. B
Fig. 18G—H

Description

Tricolporate, amb circular and shape ellipsoidal. The exine is of medium
thickness and sexine and nexine are clearly differentiated. Crassinexinous. The
sexine is tectate, finely columellate and granular. The colpi are narrow and almost
reach the poles of the grain. The ora are lalongate, slit-like and of medium length
and the costae endopori are robust. Length of illustrated specimen 27 pw, width
21 w, width of ora approximately 0,5 w, length 3 wp.

Discussion

The slit-like ora is a characteristic feature of Tricolporopollenites sp. B.

Affinity
There is no information regarding the affinity of Tricolporopollenites sp. B.

Distribution

This species was infrequent at Arnot.

Tricolporopollenites spp. C and D
Species C: Fig. 18A—B; Species D: Fig. 18C—F

Description

Both species are small and angulaperturate grains. Tricolporopollenites sp. D
has a more triangular amb than Tricolporopollenites sp. C, and both are prolate.
The surface sculpturing of the two forms also differs. The colpi are narrow and
almost reach the poles. They are deeply buried in narrow clefts shaped by the
sharp inward bends in the exine at the apices of the triangular amb. The ora is
small and protrudes into this cleft producing the characteristic ‘H-shape’ seen in
equatorial view. The exine is thick and finely columellate. Length of polar axis
18-25 w, width 13-18 w, exine approximately 2 wp.

Affinity

Ferguson (1977) has described ‘H-shaped’ aperture structures in certain
genera of the Cornaceae. These structures are formed by a pore being joined to
two lateral thinnings of the endexine that run parallel to the colpus. The fossil
types described above probably possess this structure. If the uniqueness of this

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

aperture morphology is confirmed, these fossils represent the first certain, and by
far the earliest, record of cornaceous pollen to date. The particular genus
concerned is Cornus and the Cornus sanguinea-subtype (Ferguson 1977: 6, figs 31,
4f-g, 5a-g). The fossil forms are unlike the pollen of Curtisia, the extant
monotypic southern African genus of the Cornaceae. Ferguson (pers. comm.
12 July 1984) notes that similar endoapertures also occur in at least one genus of
the Rubiaceae (Lewis 1965) and in various genera of the Escalloniaceae and
Penthoraceae, and in the genus Sedum of the Crassulaceae. The occurrence of
this feature is discussed in Hideux & Ferguson (1976).

Distribution

Both species are rare at Arnot.

Pollen with more than three apertures
Genus Retistephanocolpites Leidelmeyer, 1966

See Saxena (1982) for a discussion of this genus.

Retistephanocolpites sp.
Fig. 19E-G

Description

The pollen is tetracolpate. The colpi are about two-thirds the polar axis, wide
open at the equator and their margins are entire. The exine is relatively thick,
crassisexinous and tectate. The tectum is closed and columellate, and under the
SEM it can be seen that the surface sculpturing consists of a dense mat of fine
strands. The amb is circular. Diameter 16-20 pw, exine 1,5 w.

Discussion

See Saxena (1982) for a detailed discussion of the taxonomy of Tertiary
polycolpate forms. Many of these forms are known especially from the Indian
Tertiary record. As Saxena points out, exinal thickenings and precise sculptural
details are conservative features that are of the most use in attributing generic
status to fossil forms and suggesting their affinities. The unusual surface
morphology of the present form should lead to eventual positive identification.

Affinity
No positive suggestions can be made regarding the affinity of Retistephano-

colpites sp. The pollen of Rubia (Rubiaceae) and Catostemma (Bombaceae),
amongst others, appear superficially similar to this fossil species.

Distribution

This form is rare in levels sampled at Arnot but it is the dominant angiosperm
pollen in earlier (early Palaeocene?) sediments from Namaqualand.

PALYNOLOGY OF THE ARNOT PIPE

ee ee es

qd

Fig. 19. A-B. Grootipollis reuningii sp. nov. C-D. Crotonipollis burdwanensis.

E-G. Retistephanocolpites sp. H. Ulmipollenites sp. 1-J. Forma G. K-L. Forma F.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Grootipollis Krutzsch, 1966
Grootipollis reuningti sp. nov.
Fig. 19A-B

Etymology

This species is named after Dr E. Reuning, the geologist whose interest in
the Arnot Pipe, Banke, led to the discovery and study of its fossiliferous
sediments.

Description

The pollen is spherical and always invaginated to some degree. It is
periporate and has between fourteen and twenty pores. Triangular-shaped
verrucae of medium height, with rounded tops, are arranged in circular ‘croton
_ patterns’. Five to eight verrucae, each with an apex pointing towards the middle,
form a single circle, but this pattern is disrupted where a pore is situated. The
verrucae are quite small structures so that their triangular shape may not readily
be noticed in plan view. The exine is thick. Equatorial diameter 45—50 w, exine
approximately 3, size of verrucae (plan view) 1m, diameter of pore
approximately 3 w.

Discussion

Grootipollis reuningii is twice the size of the type species of the genus,
Grootipollis cretacius (Jansonius & Hills 1976: 1191), which also has relatively
smaller and fewer pores (8-10 in G. cretacius versus 14—20 in G. reuningii). Apart
from the obvious differences in aperture morphology, the smaller verrucae and
circular arrangement distinguish G. reuningii from the two other forms displaying
the ‘croton pattern’—Crototricolpites densus (see p. 67) and Crotonipollis
burdwanensis (see p. 80), which occur at Arnot.

Affinity

Grootipollis reuningii shows affinity with genera of the Thymelaeaceae (such
as Phaleria, Passerina and Struthiola) and to the Buxaceae (Muller 1981: 48). The
former is considered the more likely.

Distribution

Grootipollis reuningii is common at Arnot. The genus and related forms are
known from the late Cretaceous of the Northern Hemisphere (Muller 1981: 48),
but have not been recorded from Australian late Cretaceous or Tertiary
sediments.

Genus Ulmipollenites Wolff, 1934

See Srivastava (1969) for discussion on this genus.

PALYNOLOGY OF THE ARNOT PIPE 79

Ulmipollenites sp.

Fig. 19H

Description

Medium-sized, 5-colpate, aspidote form. The exine is relatively thick and the
aspides, around the short narrow colpi, appear as prominent knobs. The exine
appears undifferentiated under the light microscope and the surface sculpturing is
undulating to rugose. Diameter of illustrated specimen 24 yw, exine 1,5—2 w, exine
at aspides 3 yw, colpi width 0,75 yu.

Discussion

The present form differs slightly from available descriptions of other fossil
forms placed in the genus U/mipollenites and the related genus Ul/moidipites in the
degree of narrowness of the colpus pore and the degree of thickening of the
aspides. In other respects it is similar to extant species of Ulmus such as U. glabra
(Nilsson et al. 1977: 108). However, Ulmipollenites sp. is also similar to the
species Haloragis haloragoides (Cookson & Pike, 1953), suggesting a possible
affinity to Haloragis (Haloragidaceae) (H. A. Martin 1973: 21). More specimens
will have to be studied before the correct affinity of this form can be determined.

Affinity
The affinities of Ulmipollenites sp. are thought to be with the genus U/mus of
the Ulmaceae or with members of the Haloragidaceae.

Distribution

This species is rare at Arnot. Ulmus-like pollen appears over a widespread
area including Africa, North and South America and India during the
Maastrichtian (Muller 1981: 19; Salard-Cheboldaeff 1981). In West Africa the
form has a continuous record up to the Miocene. Haloragis- or Haloragicidites-
types are known from the Eocene of Eurasia (Muller 1981) and the Miocene in
Australia (H. A. Martin 1973).

Inaperturate pollen

Genus Crotonipollis Baksi, Deb & Siddhanta, 1979

The above name is used here despite the fact that Baksi et al. (1979)
instituted their genus apparently unaware of a prior homonymous generic
diagnosis with a very different content (De Lima 1976).

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979
Fig. 19C-D
Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979: 233, fig. 1.

Description

The grain is large, robust and inaperturate and the sexine bears large
regular-shaped, triangular verrucae arranged in the characteristic ‘croton
pattern’. Six triangular verrucae, each with an apex pointing towards the middle,
constitute a unit of the ‘croton pattern’. The grain is so robust that under the light
microscope and normal processing no details of the nexine can be distinguished.
Diameter 50 w.

Discussion

Baksi et al. (1979) discuss the differences between two Indian species of
Crotonipollis.

Affinity

Baksi et al. (1979) mention the similarity between Crotonipollis and the
pollen of extant species of Jatropha (Euphorbiaceae). A large number of genera
of the subfamily Crotonoideae are listed by Punt (1962) as possessing the
inaperturate ‘croton’-type pollen, so that determination of closer affinities within
this group must await further work. At least some species of Jatropha have
prominently ‘ribbed’ verrucae (Bonnefille & Riollet 1980: 69, pl. 52) and thus
clearly differ from the present forms. The genus Croton is a more likely
affinity—cf. Croton draco (Lonzano-Garcia 1979: 318, pl. 12).

Distribution

Crotonipollis burdwanensis is rare at Arnot. It also has rare and restricted
occurrence in the Eocene and Palaeocene of India.

Pollen found in obligate tetrads
Genus Ericipites Wodehouse, 1933
Ericipites sp. A
Fig. 20A—C

Description

Tetrahedral tetrads of tricolporate pollen with hamulate to weakly rugulate
sculpture. The amb of each monad is subcircular. The exine is relatively thin.
Apertures arranged according to Fischer’s rule (Erdtman 1952: 14); the colpi are
long thin slits, which broaden equatorially to enclose the ora. Ora lalongate. The
costae endopori are prominent. Colpi are three-quarters to two-thirds the length
of polar axis. Diameter of tetrad 24-35 mw, exine 1-1,5 wp.

81

PALYNOLOGY OF THE ARNOT PIPE

G. Dicotetradites sp.

Fig. 20. A-C. Ericipites sp. A. D-E. Ericipites sp. B. F-

H-M. Triporotetradites sphericus sp. nov. N. Dicotetradites sp.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ericipites sp. B
Fig. 20D-E

Description

Tetrahedral tetrads of tricolporate pollen. The colpi are thin slits and
relatively short and the ora are inconspicuous. Apertures are arranged according
to Fischer’s rule (Erdtman 1952: 14). The amb is subcircular to triangular with
broadly rounded apices. The nexine is robust and the sexine is fossulate, being
traversed by fine cracks.

Affinity

As Martin (1978) and others have pointed out, the tetrads of the Ericaceae,
Empetraceae, and Epacridaceae can generally not be distinguished. The extant
families are also not well known palynologically. Because of the distribution of
the modern families it is likely that the present forms have affinity to the
Ericaceae and, less likely, to the Epacridaceae.

Distribution

The earliest record of tetrads with Ericipites-like morphology may be from
marine middle Cretaceous sediments off North Africa (Kotova 1978). Forms
designated Ervicipites are first recorded in the Northern Hemisphere in European
Maastrichtian sediments and may have affinity to the Ericaceae or Empetraceae
(Muller 1981: 41). By the Eocene they can be a common element in assemblages
from central Europe (Muller 1981). An Ericipites form, Ericipites scabratus
(Harris, 1965), comparable to Evicipites sp. A above, is an infrequent element in
middle to late Palaeocene sediments from south-eastern Australia (Harris 1965)
and a further form, Ericipites crassiexinous, is common in middle to upper
Eocene strata. Ervicipites was not recorded in Maastrichtian sediments off
southern Africa (McLachlan & Pieterse 1978) nor, for some peculiar reason, was
it observed in the Neogene (?) Knysna lignites (Thiergart et al. 1963). It is a
component of the late Neogene record in the south-western Cape (Coetzee 1981).

Ericipites sp. A is relatively common at Arnot while Ericipites sp. B is rare.
This is the earliest record from the African subcontinent of a plant group that is a
prominent member of the Capensis Flora today.

Genus Dicotetradites Couper, 1953
Dicotetradites sp.
Fig. 20F—G, N
Compare

Dicotetradites clavatus Couper, 1953: 63, pl. 8 (fig. 125).
Paripollis ochesis Partridge, in Stover & Partridge, 1973: 274, pl. 28 (fig. 2).

PALYNOLOGY OF THE ARNOT PIPE 83

Description

The pollen occurs in obligate tetrads. It could not be ascertained whether the
monads are tricolpate or tricolporate, but the colpi are long and arranged in the
normal manner according to Fischer’s rule (Erdtman 1952: 14). The ora, if indeed
present, are inconspicuous and opposite one another. The robust clavae
contribute towards the difficulty of observing the ora. Each monad is subspherical
to subtriangular; the exine is thick and clearly differentiated into a nexine and
sexine. The nexine broadens to end in a knob at the margins of the colpus. The
sexine is decorated with robust verrucae or clavae which are angular in plan and
have rounded tops. These structural elements are most robust in the distal polar
region and become smaller in the equatorial region. The sexine is hardly, if at all,
present on the interfacial proximal face of each monad. Overall size very variable.
Diameter of figured specimen 45 yw, diameter of monads 30-33 yw, exine 6 pw in
the distal polar area, 3 yw in the equatorial area.

Discussion

Only five specimens were observed and the overall size range and variation in
sculpturing suggest that more than one species is present. Figure 20N shows a
grain tending towards the clavate-baculate condition described as Dicotetradites
clavatus Couper, 1953, while the well-preserved grain illustrated in Figure 20F—G
is very similar to Paripollis ochesis Partridge, in Stover & Partridge, 1973.
Because of the uncertainty as to whether the present form has an ora or not, the
original generic diagnosis Dicotetradites was provisionally preferred and the
revised diagnosis of Crosbie & Clowes (1980) was ignored.

Affinity

The affinity of Dicotetradites sp. is probably with the Epacridaceae, which
also have obligate tetrad forms with individual grains possessing verrucate
sculpturing that may obscure their apertures, and with the sexine confined to their
distal walls and absent from the contiguous proximal walls (Mathews 1966: 464,
469, pl. 2 (fig. 3)—cf. Epacris heteronema).

Distribution

Dicotetradites sp. is very rare at Arnot. The genus Dicotetradites is a common
form in the Eocene of New Zealand with a range from the Palaeocene to the late
Oligocene (Crosbie & Clowes 1980: 460). It is also known from the Oligocene in
south-eastern Australian sediments (Stover & Partridge 1973). Similar forms
have not been reported from the Northern Hemisphere or tropical Africa. Today
the Epacridaceae are found mainly in Australia and Tasmania, but also in South
America (Willis 1966).

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Triporotetradites van Hoeken-Klinkenberg, 1964
Triporotetradites sphericus sp. nov.

Fig. 20H-M

Etymology

This specific name refers to the spherical shape of the tetrad.

Description

The structure of the grain is extraordinary. It is a tetrad with each monad so
shaped that the whole grain is spherical. It could be described as a spherical, cross
tetrad with all four monads meeting at the centre. Each monad is triporate, the
pores being circular and arranged according to Garside’s rule, i.e. three pores
grouped together at four points on the surface of the grain (Erdtman 1952: 14).
The wall of each monad is psilate and is not differentiated under the light
microscope; the pore structure in this layer consists of a low, narrow but distinct
ring on the exterior of the grain, which forms the top rim of an elongated chimney
extending into the interior of the monad. Each group of pores is situated in the
hollow beneath the sexinal (?) layer formed by the curving of three adjacent
monads away from the circumference of the tetrad. The rims of the three adjacent
pores may touch one another. The four monads are enveloped by a reticulate,
simplicolumellate, undifferentiated sexinal (?) layer to form a single inaperturate
spherical grain. The reticulum is regular and perfect and the sexine may thicken
over the groups of pores. The size of the grains is very uniform, the diameter of all
measured specimens being between 31 and 34 yw, sexine varies between 1,5 and
3,5 w, diameter of pore opening 1,5-3 pw, height of whole pore structure 4—5 p,
size of lumina approximately 1 wp.

Discussion

Triporotetradites sphericus can be distinguished from Bysmapollis emaciatus
Partridge, in Stover & Partridge, 1973 (p. 273, pl. 28 (fig. 1)) in pore and exine
structure; the latter also has pores arranged according to Garside’s rule (Erdtman
1952: 14) The inaperturate, enveloping, reticulate layer of T. sphericus is the most
distinguishing feature. The only other tetrad that has pores arranged similarly is
Ajatipollis tetraedralis (Bolkhovitina) Krutzsch, 1970. In this form, however, the
pore placement is described as free, and the pores are clearly not as closely
grouped as in the present form. Crosbie & Clowes (1980: 460, figs 4, 6) note that
tetrads of the species Dicotetradites clavatus Couper, 1953, have a granulate
sexinal layer which is continuous over the junction of individual grains.

Figure 20L—M shows a grain with some of the reticulum missing and with the
psilate nexine (?) of the individual grains, as well as their pore structure, exposed.
It is likely that a new genus will eventually have to be erected to contain this form.

PALYNOLOGY OF THE ARNOT PIPE 85

Affinity

It has been suggested that members of the genus Triporotetradites have
affinity with the extant genus Gardenia of the Rubiaceae. Triporotetradites
sphericus, however, differs widely from any known Triporotetradites.

Distribution

The present form is common at Arnot. The genus 7riporotetradites is known
from the upper Eocene of Europe and the lower Miocene of the Cameroons
(Muller 1981). Muller regards a Maastrichtian record of the genus from Nigeria
(Van Hoeken-Klinkenberg 1964) as not acceptable. Stover & Evans (1973: 58)
mention an undescribed Triporopollenites type and illustrate a ‘planar tetrad’ that
superficially resembles the present form. These forms occur in late Cretaceous
and Palaeocene sediments of the Gippsland Basin, south-eastern Australia.

Dicotyledonous pollens not assigned to genus
Forma F
Fig. 19K-L

Description

Large, thin-exined 5-colpate form with a circular amb. The colpi gape
equatorially and have characteristically rounded ends. Nexine and sexine are
differentiated and the exine is subpsilate. Diameter of illustrated specimen 34 wp.

Affinity
The affinity of Forma F is perhaps with the Labiatae.

Distribution

Forma F is rare at Banke.

Forma G
Fig. 19I-J

Description

Tricolporate, very rounded triangular, planaperturate, crassiexinous, psilate
grain. Crassiendexinous. The ektexine stops short of the colpus as the endexine
thickens to form a low costae colpus. Regarding the intectate sexine the grain is
syncolpate. The endocolpus is lalongate. Equatorial diameter 25 mw, exine 2,5 wu,
endexinous costae colpus 4 w thick.

Discussion

Forma Gis a rather unusual grain with no comparable fossil forms known. In
the very thick exine and syncolpate state there is some similarity with the
following form.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity
The affinity of Forma G is unknown.

Distribution

Only one well-preserved grain was observed at Arnot.

Forma H
Fig. 13M—O

Description

The grain is tricolpate and crassiexinous. A thin ektexine may be present but
can hardly be differentiated from a massive endexine. At one pole the grain is
apparently syncolpate, the exine thinning towards this pole. There is some
irregularity in the margins of the colpus and a suggestion of a hexaporate
condition with the two pores of each colpus situated non-symmetrically around
the equator. The amb is lobate and the shape is oblate but with a clear
irregularity, the exine thinning towards the syncolpate pole. Equatorial diameter
of illustrated specimen 29 yw, exine 4 w, colpus 2 wu wide, polar axis 22 p.

Discussion

Forma H is a highly unusual grain especially in its lack of symmetry.

Affinity

There would seem to be little resemblance between Forma H and any extant
family except perhaps (but excluding the lack of symmetry) to the Gyrostemon-
aceae (Erdtman 1952: 198).

Distribution

Only one well-preserved grain was observed.

Forma I
Fig. 21A-B

Description

Only two specimens were observed and the following description is
preliminary. A medium-sized, probably tricolporate grain with an open reticulate
sculpturing developing into longitudinally striate sculpturing in the area of the
colpi fossae.

Discussion

As far as is known, no very similar fossil forms are known. The reticulate
sculpturing illustrated in Figure 21B resembles Alangiopollis eocaenicus as
illustrated in Reitsma (1970: 283, pl. 33).

PALYNOLOGY OF THE ARNOT PIPE

F

Fig. 21. A-B. Formal. C-F. Fenestriorites (photomicrographs not of material from
Arnot).

87

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

The affinity of Forma I is possibly with the Alangium kurzii-type of the
Alangiaceae, Section Marlea, as described by Reitsma (1970). If this is correct
then, although Forma I is a new specific record, the pattern that all early fossil
occurrences of this family are of the primitive Section Marlea is maintained.

Distribution

Forma I is rare at Arnot. Alangium kurzii and A. rotundifolium are found
today in forests of Indo-China.

POLLEN COUNTS

The pollen diagram (Fig. 3A) expresses the value of individual elements as
percentages of the total number of grains counted for each of the seven sampled
levels. Only values of one per cent and greater are included.

Little can be deduced from the pattern revealed by the diagram. Triorites
operculatus is always the most common form. A sharp increase in its relative
abundance occurs towards the top of the diagram where it achieves values greater
than 40 per cent. This increase does not correlate with simultaneous changes in
abundance of any other elements, but is preceded by a sharp increase in the
abundance of Stereisporites (Sphagnum) and the other spores. These last two
elements clearly covary. A likely explanation for this peak in their abundance is
that some change in the local geography allowed greater run-off and water trans-
port to the depositional environment. The sample from 20-21 m (65-70 feet) is
not highly carbonaceous so that the immediate presence of a peat bog is not
indicated. The increase of Stereisporites together with the other spores (which
include two possible tree-ferns— Cyathidites and a member of the Lophosori-
aceae) suggests that all grew in the same environment. This may have been a
forest with the plants concerned growing either on the floor (moss) or as part of
the understorey (tree-ferns and other ferns).

The occasional peaks in the abundance of Clavatipollenites, Triorites
arnotiensis and Tricolporopollenites brinkiae may represent changes in local
edaphic conditions and/or a slight shift in a vegetation boundary. Some extant
genera of Chloranthaceae (represented at Arnot by Clavatipollenites) are forest-
margin species. The coherent patterning in the percentage values of Triorites
operculatus and Dicolpopollis sp. are possibly indicative of long-term change in
the regional vegetation and there is some basis for speculation on its nature (see
point 21 of the discussion).

The percentage values for conifers varied between 15 and 5 per cent.

DISCUSSION

The body of this research has consisted of systematic, descriptive palynology.
The affinity of as many fossil forms as was possible has been noted and the results
of this work are summarized in point form below. The detail achieved allows for

PALYNOLOGY OF THE ARNOT PIPE 89

some comparisons to be drawn with the Australian and tropical African early
Tertiary palynomorph records, and some statistics to be produced on the
taxonomic levels and degree of extinction that has occurred in Africa between the
early Tertiary and the present.

The described assemblages from Arnot are isolated in time and space.
Although subsequent work has provided more biostratigraphic evidence, long
continuous sequences and a stable dating framework are not yet available. The
fact that only seven samples from a short 25 m sequence could be analysed makes
it difficult to evaluate the changes recorded.

The small amount of work done on palynomorph assemblages from
kimberlite pipes from the northern Cape and from Botswana and the general
paucity of published work on local late Cretaceous and Tertiary palynology make
discussion on vegetation history in these time ranges on the subcontinent
premature. However, the evidence produced in this study provides a much better
base than was previously available for tentative statements regarding the
vegetation represented at Arnot and the palaeoclimate involved. Furthermore,
since other authors (Axelrod & Raven 1978; Tankard & Rogers 1978) have
placed their interpretations upon the previously published palaeobotanical
evidence from Arnot, it was thought that some remarks on these subjects were
necessary.

Some of these remarks rely upon suggested features of the local and regional
geography of the site and its mechanism as a pollen trap for their support. These
features (see p. 5) may be summarized as follows:

The regional topography was that of a relatively mature, flat landscape
unassociated with any prominent montane region. The country rock of the area is
Namaqualand gneiss, which was blanketed by base rich ‘kimberlitic’ material in
the vicinity of the Arnot Pipe. The cones of ejectamenta of the numerous
‘kimberlite’ volcanoes are thought to have been relatively low (perhaps in the
order of 100—150 m) and the infilling of each crater to have been completed a few
million years (maximum) after emplacement. The crater lake formed in the vent
of the palaeo-Arnot Pipe volcano accumulated laminated, fine-grained, carbon-
aceous shale and mudstone sediments towards its centre and acted as a local,
small basin pollen trap, unrelated to developed drainage patterns. The agents of
pollen transport would have been wind, local run-off, and probably settling out of
suspension of fine sediments and pollen in the quiet centre of the lake after
gravity avalanching of material off the talus slope of the cone.

Although the slopes of the volcanic cones must have provided special
edaphic conditions favouring certain plant species, these areas were isolated
features in the general landscape. Even the plants that occurred on the local
edaphic site would have formed part of a more generalized, wider distribution of
plant associations. There is no climatological reason to suppose that the Palaeo-
cene location of the region experienced a transition zone between climatic regimes.

In terms of the present knowledge available, therefore, it seems safe to
suggest that the species of plants recorded at Arnot grew on base rich soils and

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

were part of a widespread non-montane Palaeocene vegetation growing in the
interior of the subcontinent.

1. The following plant familes are represented in the Arnot palynoflora:
(a) Pteridophyta and Bryophyta: Sphagnaceae, Cyatheaceae, Anthocerotaceae,
Polypodiaceae, Schizaeaceae and Ophioglossaceae.
(b) Conifers: Podocarpaceae and Araucariaceae.
(c) Angiosperms: Chloranthaceae, Restionaceae, Palmae, Ulmaceae (UI-
moideae) or Haloragidaceae (?), Casuarinaceae or Myricaceae, Proteaceae,
Gunneraceae, Euphorbiaceae, Thymelaeaceae, Anacardiaceae, Cornaceae, Eri-
caceae, Epacridaceae and Caesalpinaceae.

Several more tentative suggestions about the possible affinity of fossil
morphotypes to modern families are also made in the text.

2. The lack of diversity in the families Proteaceae, Ericaceae and Res-
‘tionaceae is notable. These three families are at present large and prominent
components of Cape fynbos vegetation. By Eocene times the Proteaeceae are
highly diversified in Australia (Martin 1981) and are represented by many forms
in the Knysna lignites (Thiergart et al. 1963), which may be either Eocene—
Oligocene (Thiergart et al. 1963; Helgren & Butzer 1977), or early Neogene in
age. Both subfamilies of the Proteaceae, the Persoonioideae and Grevilleoideae,
may be represented in the Arnot assemblages but no proteaceous form is
common to the Arnot and Australian early Tertiary assemblages. Most of the
proteaceous forms probably represent extinct genera, but the genera Leucosper-
mum and Protea may be represented. The evidence from the pollen morphology
of the Restionaceae suggests that present ideas about evolution within this family
may need revision (see Johnson & Briggs 1981).

3. The diversity in the Euphorbiaceae (non-heathland types, Specht 1981:
790) is notable. The evidence for early presence in the African subcontinent of
species with affinity to the Euphorbiaceae, Thymelaeaceae, Monimiaceae,
Anacardiaceae, Rubiaceae (?) and Ulmaceae (?), amongst others, is also
important and provides valuable data for a perspective on the in situ evolution of
plant phylogenies in the subcontinent. Wood of Euphorbiaceae and Monimiaceae
has been identified in late Cretaceous deposits on the east coast of southern
Africa (Miiller-Stoll & Madel 1962).

4. In terms of biostratigraphic age-bracketing, none of the evidence
contradicts a possible Palaeocene date for the sequence. The assemblage is
composed of a mixture of: (a) forms whose affinity to modern families can be
traced, and (b) archaic forms known mainly from, or with a record extending
back into, the Cretaceous. This combination suggests proximity to the Cre-
taceous—Tertiary boundary (see points 5 and 6 below). On the other hand
Monoporites annulatus (Gramineae) while prominent in tropical African Eocene
assemblages (Salard-Cheboldaeff 1979, 1981) does not occur at Arnot. This may
support a pre-Eocene age for these assemblages.

PALYNOLOGY OF THE ARNOT PIPE 91

5. The following form-genera and species present at Arnot are known
mainly from the Cretaceous:

Araucariacites sp., Zonalapollenites, Monocolpopollenites, Fenestriorites, Ali-
sporites grandis, Foveotriletes margaritae, Cicatricososporites, Distaverrusporites
and Hamulatisporis.

In addition Arecipites, Liliacidites, Tricolpites reticulatus and Clavatipollen-
ites forms are common in the late Cretaceous but continue through the Tertiary,
and Araucariacites australis exits from the tropical African record in the late
Cretaceous and from the Indian record at the Cretaceous—Tertiary boundary, but
has a continuous record into the Tertiary in Australia.

6. Comparing the Arnot palynomorph assemblage with other Palaeocene
assemblages* and with extant floras produces the following rough estimates:
(a) 57 per cent of the species are unique to the Palaeocene of the African
subcontinent;
(6) 59 per cent no longer occur in Africa;
(c) 39 per cent are extinct;
(d) 39 per cent of the forms, and
(e) 60 per cent of the families, are common to the Arnot and south-eastern
Australian Palaeocene, while only
(f) 12 per cent of the forms, but
(g) 65 per cent of the families, are common to the Arnot and tropical African
Palaeocene.
(a—d excluding pteridophytes, no families of which are known to have become
extinct during the Cenozoic.)

These statistics are employed in the discussion that follows.

7. At generic levels (refer to 6e above) the Arnot Palaeocene flora is more
closely related to south-eastern Australian than to tropical African Palaeocene
floras. This reflects in the main a common southern Gondwana pteridophyte and
conifer floral inheritance. However, a degree of similarity in the climates of the
two regions is also indicated. The low level of commonality at generic rank (6a, f)
between the Arnot and tropical African assemblage suggests a marked difference
in the climates experienced in the two regions. This pattern is superimposed upon
a continuing phytogeographical relationship expressed at a higher taxonomic
level (see 6g). The uniqueness (see 6a) of the Arnot flora mainly emphasizes its
difference from the tropical African Palaeocene flora.

Points 6b and 6c are measures of the antiquity of Palaeocene floras and
although comparable estimates are not available, a rough comparison does
suggest that the level of ‘modernity’ encountered in south-eastern Australian
Palaeocene flora might be similar to that of the Arnot flora and dissimilar to the
state of the tropical African Palaeocene flora. It would appear from the work of
Salard-Cheboldaeff (1978, 1979, 1981) that this latter flora contains many archaic

* The studies on which these comparisons are based include Germeraad et al. (1968), Salard-

Cheboldaeff (1978, 1979, 1981), Martin (1978, 1981), Harris (1965), Kemp & Harris (1977),
Kemp (1981), Stover & Partridge (1973), and Muller (1981).

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

forms and that, in the equatorial region, continuity with Neogene floras is only
marked from the Upper Eocene—Oligocene (Salard-Cheboldaeff 1981: 435).

The difference of 20 per cent between 6b and 6c is a measure of the level of
extinction that has occurred in Africa between the Palaeocene and the present. It
suggests the contrast in this continent between the Palaeocene environment—
with perhaps more equable climates and easy transitions between climates—and
the subsequent global development of more distinct zones of climate and
vegetation, some of which were not well represented in Africa. In particular,
Africa does not extend into high southern latitudes and therefore lacks an
extensive zone of temperate climate and the role that this might play on a
continental scale in the evolution of vegetation. For this reason, in addition to the
probable relative aridity of Africa in the Cenozoic, the subtropical—temperate
vegetation of Africa has suffered a relatively high degree of extinction between
the Palaeocene and the present.

8. The connections and contrasts of the Arnot palynoflora with early

Tertiary south-eastern Australian floras can be further elucidated.
(a) The low percentage representation of spores at Arnot relative to the
Australian norm (5% versus 20-30%) may be explained either by reference to
the general pattern of low spore representation in Africa (Salard-Cheboldaeff
1981) or by the peculiar geographic setting of the Arnot site. Relevant to the
former explanation are the suggestions that Africa has been relatively drier than
Australia for a long time and/or that the pattern reflects the intracontinental
location of the sites producing this pattern, i.e. east coast of Australia, west coast
and interior of Africa.

The latter explanation suggests that whereas it can be assumed that most
spores reach a depositional environment via water transport, this agent of
transport (as already indicated, see p. 88) may have played a minor role in the
geographical setting of the Arnot Pipe. Although involving an obviously circular
argument, there is some support in the covariation of values for Stereisporites and
the other spores for the suggestion that an increase in water transport explains the
single anomalously high value of 20 per cent recorded for these two elements at
the 65-70 foot (20-21 m) level (see Fig. 3).

(b) Apart from relatively high spore representation (indicating high humidity
and/or equability), the south-eastern Australian conifer flora is dominated by
podocarpaceous taxa, especially the genera Dacrydium and Microcachrys. In
contrast Araucariaceae entirely dominate earlier Palaeocene or late Cretaceous
assemblages from the Arnot region and make up about half of the conifer
representation at Arnot. This contrast is explained by the comparatively high
latitude of south-eastern Australia in the Palaeocene (65°S versus 40°S today) and
the correspondingly lower temperatures and more temperate climate experienced
there.

However, Araucariacites is hardly represented at all at a more northerly and
inland Australian site situated at about 50°S in the same time range (Wopfner
et al. 1974).

PALYNOLOGY OF THE ARNOT PIPE 93

Although the non-occurrence of a form at a particular site is not a reliable
observation, it is to be noted in comparing the Palaeocene Australian conifer flora
with the Arnot flora that Microcachryidites, Dacrycarpus, Dacrydium franklinii-
type, and Phyllocladus are not recorded at Arnot. All of these forms, however,
except the Dacrydium franklinii-type and the Dacrydium cupressinum-type
(recorded at Arnot) are known to occur in the late Cretaceous in the south-
western Cape (McLachlan & Pieterse 1978). Their non-occurrence at Arnot may
indicate the existence of a marked climatic gradient between the palaeolatitudes
of Arnot and the south-western Cape or it may simply indicate the existence of a
montane habitat in the latter region. It is clear, however, from evidence from
elsewhere (Herngreen & Chlonova 1981: 506, 511) that Microcachryidites is a
sensitive indicator of some climatic gradient.

Microcachryidites is still a prominent component of the Neogene vegetation
of the south-western Cape, while Araucariacites does not appear in the Neogene
record (Coetzee 1978a, 1978b, and pers. comm.). No species of Microcachrys
occur today in Africa.

It would appear that one Zonalapollenites form as well as Podocarpidites
riembreekensis and perhaps Podocarpidites kamiesbergensis are at present unique
to the southern African Palaeocene.

(c) The absence of a number of angiospermous forms characteristic of the
Australian Palaeocene from the Arnot assemblages provides further contrasts.
These include the three Nothofagus pollen types (although these are only
common from the Eocene in Australia), Myrtaceae, Olacaceae (Anacalosa),
Euphorbiaceae (Austrobuxus—Dissiliaria), Banksieae, Xylomelum-type and
other extinct forms attributed to the Proteaceae, Santalaceae, /lex (Aquifoli-
aceae) and Cupanieae (Sapindaceae).

On the other hand the following forms that occur at Arnot are not known
from the Australian Palaeocene: Triorites operculatus and T. sphericus (the
former very prominent), Retistephanocolpites (prominent earlier (?) in the
Palaeocene from other sites in Namaqualand), Crototricolpites and Crotonipollis
(Euphorbiaceae), Grootipollis (Thymelaeaceae), all the proteaceous forms
(3-4?), Milfordia (Restionaceae), Rhus (Anacardiaceae), Triporotetradites,
two monocolpate forms (one, and perhaps both, of which have affinity to
the Palmae), a palmaceous Liliacidites, Tricolporopollenites arnotiensis and
T. brinkiae (Rubiaceae, Euphorbiaceae?), and Tricolporopollenites spp. C & D
(Cornaceae).

9. Apart from the absence in the tropical African late Cretaceous—early
Tertiary record of the spore and conifer ‘temperate’ southern Gondwana floral
component, the most important contrast between the Arnot and tropical
palaeofioras lies in the representation of palms. In the Palaeocene of tropical
Africa, twelve palm form-genera are recognized and their representation is
constantly high (20-25%) (Salard-Cheboldaeff 1981). At Arnot two to three
forms with affinity to the Palmae occur and their contribution to the palynomorph
assemblages never rises above 3 per cent.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

As already indicated very few form-genera occur both at Arnot and in
Palaeocene tropical African palynomorph assemblages (12 %), while the relation-
ship is closer at a high taxonomic level, with 65 per cent of the families being
common to both.

The forms common to both regions are Foveotriletes margaritae, Distaverru-
sporis, Zonalapollenites sp. B (Cingulatipollenites), Monocolpopollenites sp. B,
Celtidoideae? (Triorites operculatus—Triorites festatus/tenuiexinus), Crototricol-
pites, Ulmipollenites sp., Proteaceae (Propylipollis meyeri—Proteacidites de-
haanii).

The families Olacaceae, Ctenolophonaceae, Malphigiaceae, Moraceae,
Acanthaceae, Mimosaceae, Bombaceae, Apocynaceae, Balanophoraceae, Mela-
stomaceae, and Combretaceae are recorded in tropical African assemblages (with
some genera being prominent) but are not known from Arnot.

A less biased reflection of differences between the tropical African and
- Arnot Palaeocene floras is provided by the list of families that, as far as is known,
occur only in the latter region: Anthocerotaceae, Sphagnaceae, Podocarpaceae,
Araucariaceae, Chloranthaceae, Myricaceae, Ericaceae, Epacridaceae, Gunner-
aceae and Cornaceae.

10. These comparisons introduce an attempt to reconstruct the palaeo-
vegetation and climate of the Arnot region. Two informal methods are used to
achieve this end. The first (points 15-18 below) involves bracketing the Arnot
climate between what has been suggested were the conditions pertaining to
palaeolatitudes to the north, south, and perhaps east of it. The second (points
18-19 below) involves extrapolating from knowledge of the present-day
distribution, habitat requirements and common growth forms of a set of taxa, to
form a hopefully coherent picture of the palaeovegetation. Table 2 provides a list
of the families and genera thought to be represented in the Arnot palynoflora,
together with notes on their present distribution, habitat and common growth
form.

11. The limitations of the second method are well known. It is clear, merely
from the presence of both Araucariacites and the Dacrydium cupressinum pollen
type at Arnot (compare the present-day habitat requirements of the genera given
in Table 2), that a process of differentiation of climates and vegetation has
probably occurred between the Palaeocene and today. Taxa such as Araucaria
and Dacrydium must have evolved relatively narrower habitat tolerances through
time, i.e. are confined to those habitats most suited to their evolutionary
potential, as a response to the development of a greater range and distinctiveness
of climates through time (cf. Kemp 1981: 40). The suggestion is therefore that
both the vegetation and climate recorded at Arnot were part of more uniform,
equable and extensive distributions of climate and vegetation (see point 13
below).

In this connection it is relevant to note that Truswell & Harris (1982: 71) in
their review of the palynology of the Eocene of Australia also mention the
occurrence of palynological assemblages containing a mixture of elements, which

PALYNOLOGY OF THE ARNOT PIPE 95

are not found growing together today, e.g. tropical rain forest and temperate rain
forest taxa.

12. Along with this observation goes the realization that it is only in a
qualified sense that concepts such as ‘tropical’ or ‘warm temperate’ can, by
extrapolation from the present, be used to describe the Palaeocene vegetation or
climates. This is not only because Palaeocene climates may have been very
different from modern climates, but also because these concepts are not basic
enough categories with which to understand even modern-day vegetation of
climatic types and distributions. Webb & Tracy (1981), for instance, use twenty-
one categories to describe the range of Australian rain-forest structural types and
nine categories to link these structural types to climatic and edaphic factors. They
argue that the recognition of the structural types enable better phytogeographical
and historical biogeographical analyses to be made between Australian and
extra-Australian regions, a claim that suggests that they have developed basic
categories with which to understand their phenomena.

13. At a general level, the mechanics of global air circulation systems are
relatively simple and therefore amenable to extrapolation to the past. Using these
mechanics and assuming the equator—pole temperature gradient of a polar-
ice-cap-free world, Lamb (1972) has modelled pre-Oligocene atmospheric
circulation. Between the equator and 60° latitude the development of pressure
anomalies would have been counteracted, not by prevailing lower and upper
atmosphere winds as is the case today, but by moving cells of higher and lower
pressures, cooler and warmer air. A weak and erratic Ferrel-type circulation
would have resulted. This type of circulation, involving the lack of zonal wind
systems, is essentially unknown today and an implication of this model is that
there are no close present-day analogues for pre-Oligocene climates and
vegetation.

This is not to suggest that, in terms of this model, some climatic gradient
between 0 and 60° latitude did not exist. An insolation gradient alone must
presumably have existed, but Lamb’s model suggests that pre-Oligocene climates
in low and middle latitudes were more uniform, perhaps more equable, extensive
and mildly transitional than is the case today. The climate of a particular region
would still have been determined by its latitude and location in terms of the
configuration of warmer and cooler seas and continents, mountains and lowlands,
and its continentality.

However, Parrish & Curtis (1982) and Parrish et al. (1982) do not accept
Lamb’s model for pre-Oligocene climates and, using an analysis of the
distribution of upwelling and organic-rich rocks and evaporites, have determined
atmospheric circulation and precipitation patterns. From this they have produced
a series of global maps portraying the broad relative isohyets of precipitation
predicted by their climatic model for various stages in the past. For the time range
concerned Parrish et al. (1982: 80) suggest that the Arnot region would have
received a relatively low to moderately low rainfall.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Visualizing the Arnot palaeovegetation and climate in relation to those to the
north and south should be done with Lamb’s (1972) model and the predictions of
Parrish et al. (1982) in mind. It is at present not clear how real the contradictions
between the two models involved are. Kemp (1978, 1981) has used Lamb’s model
to reconstruct early Tertiary Australian climates and vegetation.

14. Both the predictions of Parrish et al. (1982) and the features of
atmospheric circulation described by Lamb (1972) allow one to maintain (contra
Axelrod & Raven 1978) that there is no immediate climatological reason why,
during the time range concerned, a zone of sharply transitional climate should
have been located in the general Arnot region producing an ecotonal state of
vegetation. Also, as stated in a previous section, there is no reason to assume that
high relief existed in the Arnot region. For these reasons, and until evidence to
the contrary exists, one should attempt to reconstruct the Palaeocene Arnot

vegetation as a single unit.

15. The comparisons made between the Arnot and tropical African
palynofloras (points 6 and 9 above) highlighted the paucity of common elements.
The Arnot palaeoflora was clearly not part of a Palaeocene African ‘tropical’
flora.

However, it should be noted that Salard-Cheboldaeff (1981) has suggested
that the climate of Africa in low latitudes during the Maastrichtian was warm,
temperate and dry and that the drastic floral change recorded at the Cretaceous—
Tertiary boundary could be explained by a cooling episode in the early
Palaeocene. Many palynological studies have documented floral change at the
Cretaceous—Tertiary boundary (Muller 1980) and a Palaeocene cooling is
generally accepted to have occurred world-wide. There is disagreement,
however, as to the scale of change attributable to this factor and its relationship to
the complex global environmental changes that took place at this time (Muller
1980). Further evidence relevant to low-latitude climates in the Palaeocene comes
from Muller’s (1980) reconstruction of the latitudinal ranges of various
thermophilous taxa through the Palaeogene. This reconstruction suggests that
their ranges were most constricted in the Palaeocene, i.e. that during the
Palaeogene temperatures were lowest in the Palaeocene itself.

16. The exiting of Araucariacites from the tropical African record during the
late Cretaceous and at the Cretaceous—Tertiary boundary in India, and its
continued occurrence in the Arnot region and in Australia in the Palaeocene, can
obviously not be explained only in terms of cooler episodes in the Cretaceous and
Palaeocene (Australia and the Arnot region at mid-latitudes would at least have
had cooler winters than the equatorial regions), but must also be related to
increased precipitation or the development of more equable climates in the
tropics. The suggestion is, therefore, that in addition to a climate cooler than that
of the tropics, a climate drier or less equable than that experienced in the
Palaeocene ‘tropical zone’ is indicated by the occurrence of Avraucariacites at
Arnot.

|
:

PALYNOLOGY OF THE ARNOT PIPE 97

However, there is a possibility that factors other than climatic change could
affect the abundance of conifers relative to angiosperms in a vegetation. Doyle
et al. (1982) have recently reviewed hypotheses explaining the prominence of
gymnosperms (Cheirolepidaceae, Araucariaceae, and Podocarpaceae) in the
mid-Cretaceous tropical floras, their subsequent decline in the late Cretaceous,
and the concomitant rise to dominance of the angiosperms. Apart from the
limited evolutionary potential of conifers in the tropics, imposed by their lack of
vessels and stereotyped leaf morphology and photosynthetic ability, the use made
by their competitors—the angiosperms— of insect pollination, to produce highly
dispersed populations and high species diversity may have contributed to their
demise. “With continued diversification, the ability of angiosperms to pack more
species into a given area might have eventually led to the collapse of gymnosperm
communities by competition from many sides and dilution of populations below a
level of effective wind pollination’ (P. J. Regal, pers. comm. in Doyle et al. 1982:
86). It should be noted that, to judge from pollen morphology, a large component
of the Arnot angiosperm palaeoflora was anemophilous, perhaps leaving the
coniferous component in a better position to maintain minimum population
densities (see point 21 below).

17. The comparisons made between the Arnot palynoflora and the south-
eastern Australian record (points 6 and 8 above) suggest that the Arnot
palaeoclimate was neither as cool nor as humid or equable as the ‘temperate’
climate indicated (especially by the composition and representation of the conifer
and spore flora) for south-eastern Australia. The odd occurrence of putative
‘tropical’ indicators such as Anacalosidites or Cupanieidites in the Australian early
Tertiary (Wopfner et al. 1974: 47) may be recording the tolerance of these taxa to
low levels of insolation in combination with high equability of climate (the effect
of eastern location of sites, a warm palaeo-Pacific Ocean, and low relief?) rather
than high ‘tropical’ temperatures.

18. As might have been predicted, therefore, the comparative sandwiching
of the Arnot palaeoclimate indicates that it was a warm, moderately equable and,
relative to the ‘tropical’ norm, dryish type. Climates of this general type are found
in subtropical and warm temperate regions today. The fact that the present-day
distributions of most of the taxa listed in Table 2 are within the tropics and
subtropics therefore lends some support to the above climatic reconstruction.

The best modern analogy for the vegetation that could have grown under
such a climate might be some of the drier forest types of east Africa, or perhaps
the mixed araucarian notophyll or microphyll vine forests of north-eastern
Australia. These are described by Webb & Tracey (1981: 626, fig. 4) as moist
forest types growing under a mean annual rainfall of 700-1 200 mm.

19. An inspection of the information on growth forms in Table 2 suggests
that the vegetation in the Arnot region was forest. Trees are the most common
growth form and lianes, epiphytes, tree-ferns, forest-floor mosses and forest-
margin species are possibly also represented.

98

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Families and genera that, it is suggested, occur at Arnot, with notes on their present distribution,
size and common growth forms. (Information mainly from Willis 1966.)

Taxon

Alangiaceae (?)

(2 genera, 20 species)

Anacaridaceae

(60 genera, 600 species)
Rhus (250 species)

Anthocerotaceae

_Araucariaceae

(2 genera, 38 species)

Cactaceae (?)

(50 genera, 2 000 species)

Caesalpinaceae

Bauhinia (?) (30 species)

Casuarinaceae

(2 genera, 65 species)
Casuarina (?) (45 species)

Cornaceae

(12 genera, 100 species)

Cornus

Cyatheaceae

Epacridaceae (?)

(30 genera, 400 species)

Ericaceae

(50 genera, 1 530 species)

Euphorbiaceae

Adenocline (18 species)

or Klaineanthus (1 species)
Zimmermania (?) (4 species)
Croton (?) (750 species)

Notes

Tropics. Trees and shrubs.

Chiefly tropical, but also warm temperate areas. Trees
and shrubs.

Widespread in tropical and subtropical regions. Much-
branched shrubs, or, more rarely, trees.

Mosses mainly with a circumboreal distribution, but also
in the Mediterranean region. Hygrophytic on slightly
wet sandy soils rich in loam and mostly near forests
(Boros & Jarai-Komldédi 1975).

Southern Hemisphere, except Africa. Trees. Moist sub-
tropical and tropical non-monsoonal forests. In
Argentina Araucaria araucana is dominant in the
forest of the Subantarctic floral province.

Xerophilous growth forms of the most pronounced type.
Chiefly in the drier regions of tropical America, but
also reaching British Columbia and Patagonia. In
forest regions there are several epiphytic genera. One
genus, Rhipsalis, in Africa.

Warm regions. Mostly lianas; also trees and shrubs.

Trees or shrubs, often of weeping habit.

East Africa (? native), Mascarene Islands, Australasia.

Northern and Southern hemispheres; temperate regions
and on mountains in the tropics. Trees and shrubs,
rarely herbs.

Trees. Europe, east Asia and North America.

Tree-ferns on all southern continents.

Indo-China to New Zealand, Hawaii, South America but
chiefly Australia and Tasmania. Representing the
Ericaceae of other continents. On heaths and boggy
ground. Mostly like Ericaceae in habit, usually shrubs
or small trees.

Confined to Africa, Mediterranean and Europe in two
main masses separated by the Sahara. Cosmopolitan,
usually confined to high altitudes in the tropics; also
on moors, swamps, and peaty soils. Woody; small
undershrubs to large shrubs and a few small trees.

One of the largest plant families. Cosmopolitan in
tropical, subtropical, and warm temperate regions.
Trees in the tropics; also herbs and shrubs.

Southern Africa. Herbs.

Tropical west Africa. Giant forest trees.

Tropical east Africa. Trees.

Tropics and subtropics. Trees.

PALYNOLOGY OF THE ARNOT PIPE 99

Taxon

Gunneraceae
(1 genus, 50 species)

Lophosoriaceae
(2 genera, 2 species)

Monimiaceae
(20 genera, 150 species)

Ophioglossaceae
(4 genera, 70 species)

Palmae
(217 genera, 2 500 species)

Chamaedorea (?) (100 species)

Podocarpaceae
(6 genera, 125 species)

Dacrydium (25 species)

Polypodiaceae
(50 genera)

Proteaceae
(62 genera, 1 050 species)

Restionaceae
(28 genera, 320 species)

Rubiaceae
(500 genera, 6 000 species)
Anthospermum (?) (50 species)

Schizaeaceae
Schizaea (30 species)

Sphagnaceae
(1 genus, many species)

Thymelaeaceae
(50 genera, 500 species)

Notes

In the tropics and southern temperate regions. Perennial
thizomous herbs. Widespread in southern Africa,
except South West Africa—Namibia.

Small tree-ferns of tropical South America.

Chiefly southern tropical and especially in the ‘oceanic’
floral regions. Shrubs and trees with leathery ever-
green leaves.

Tropical and temperate regions. Small herbs, and some
tropical species are epiphytic.

Tropical and subtropical. Some are widespread, but most
genera are well localized. The palms form a charac-
teristic feature of tropical vegetation. Trees.

Warm America. Small reedy palms often forming suckers.

Southern conifers. Present on all southern land-masses.
Wide range of habitats; lowland heaths and scrubs,
open forest, rain-forest and subalpine vegetation.
Trees or shrubs.

Indo-Malaysia, Tasmania, New Zealand. Trees and
shrubs. Temperate rain-forest. Cool wet sclerophyll
forest. Mainly found in cool temperate Tasmania.
‘Frees:

Cosmopolitan, especially in the wet tropics. Almost all are
epiphytes.

Tropical Asia, Australasia, South America; tropics and
temperate areas, also mountains in Africa, South
Africa and Madagascar. The great majority live in
regions where there is annually a long dry season.
The primitive members of this family are mostly
rain-forest trees.

Mostly in southern Africa and Australia; a few in New
Zealand, Chile, Indo-China and tropical Africa.
Xerophilous, perennial with a tufted or creeping
rootstock.

One of the largest plant families. Most are tropical but a
number are temperate. Trees, shrubs and herbs.
Africa and Madagascar. Widespread, associated with
afromontane vegetation. Low shrubs and herbs.

Mainly in the tropics, but also North America. All
southern continents.

Peat-moss family. Forms peat bogs and is common on wet
forest floors and shaded mountain seeps.

Temperate and tropical regions, especially in Africa. Most
are shrubs, but there are some trees and a few lianas
and herbs.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Taxon Notes

Ulmaceae (?) Cosmopolitan. Mainly in the Northern Hemisphere tem-
(15 genera, 200 species) perate regions and tropics.
Celtis (?) (80 species) Cosmopolitan in tropics and temperate areas. Very

widespread in southern Africa, except South West
Africa—Namibia. Trees with a range of adaptability
and growth form.

Ulmus (?) (45 species) (Elm.) North and south temperate regions. Trees.

20. Can any suggestion be made as to the habitat of members of the
Ericaceae, Epacridaceae, Restionaceae and Proteaceae, and perhaps also
Rubiaceae and Thymelaeaceae, which may not have been, or were not, trees
within the suggested general forest environment? What implications does this

have for ideas about the origins and evolution of the Cape fynbos?
. No very positive statements can be made, but there are some suggestions that
these taxa could have been part of the understorey of a dryish open forest type.
The implication is that the origin and evolution of the fynbos is linked, in its
earlier stages, to the history of this vegetation type rather than to the history of
cooler, wetter, perhaps more closed-canopy forest types, such as may have been
present to the south of Arnot, or in montane situations (Coetzee et al. 1983).

The following observations provide the basis for these statements:

(a) The counter-suggestion that the distribution of these taxa may have been
controlled by the occurrence of specific edaphic conditions receives little support
from the available evidence. There is no floristic indication of the existence of
swampy or water-logged conditions in the vicinity of the site and the suggested
local topography (p. 5) argues the same. Also, the local substrates and sediments
of the pipe itself indicate that no oligotrophic soils occurred in the vicinity of the
pipe. Therefore, Specht’s (1979, 1981) hypothesis as to the possible origin of
sclerophyllous taxa cannot, in this instance, be supported. In terms of his
hypothesis, under a warm, humid, equable climate local edaphic sites such as
water-logged areas, or areas of oligotrophic soils, could have been the areas
where early sclerophyll communities originated.

(b) Ericaceae are not recorded in the Knysna lignites (Thiergart et al. 1963),
which may be either Eocene—Oligocene or early Miocene in age, and are very
rare in the early Miocene lower levels of the Noordhoek occurrence (Coetzee
1978a, 1978b). There is evidence to suggest that the vegetation types represented
at these sites were adapted to relatively wetter and more montane and equable
climates than the Arnot palaeovegetation. Restionaceae are also either very rare
or absent from the lower levels at Noordhoek, but are the most abundant type in
the Knysna lignites. These latter restionaceous forms, however, are different to
the Arnot (Milfordia) forms, having graminoid-type apertures, and their
abundance has been taken to indicate extensive marshlands in the vicinity
(Thiergart et al. 1963).

PALYNOLOGY OF THE ARNOT PIPE 101

(c) Evidence from Arnot and Botswana (Scholtz & Deacon 1982; Coetzee et al.
1983) suggests that some zonation of forest vegetation existed during the late
Cretaceous and early Tertiary in the subcontinent, and that forms with affinity to
Restionaceae, Ericaceae and Proteaceae did occur in the probably relatively drier
north-western interior.

(d) Monulcipollenites confossus, a restionaceous form, exits from the tropical
African record at the Cretaceous—Tertiary boundary (Salard-Cheboldaeff 1979).
This pattern may relate to the development of wetter and more equable climates
and closed-canopy forest during the Palaeocene. In this connection Whitmore
(1975, quoted in Webb & Tracey 1981: 613) has attributed the poverty of the
south-east Asian rain-forest grass flora to the relative stability of these humid,
closed-canopy forests. In more open and disturbable forest types (i.e. under drier
and less equable climates) the evolution of nomad grass species is favoured.

The generalized point is of importance in this discussion. Drier, less equable,
open-canopied forest types are the forest types within which an understorey will
evolve. The origins and evolution of the fynbos may therefore be linked to the
history of drier forest vegetation. In this view the fynbos shares an origin with
other generally ‘subtropical’ vegetation associations from which it has been
separated by the subsequent development of more diverse climates.

If this hypothesis is correct and it can be more adequately demonstrated that
in the early Tertiary members of the Proteaceae, Ericaceae and Restionaceae
were widespread in non-montane vegetation, then their supposedly typical
present-day distribution in Africa, high diversity in, and dominance of, the
montane vegetation of the south-western Cape, and association with nutrient-
poor soils are all ‘secondary’ features. The present pattern of their distribution
should then be viewed as being achieved as climates diversified and became, in
general (excepting especially mountains that receive orographic rain), more arid
through time. In this process earlier dominant types of vegetation and taxa were
presumably eliminated and in situ evolution of other components of vegetation
resulted in new vegetation associations, specialist adaptions, etc.

(e) The above evidence and argument suggest that Johnson & Briggs’s (1981:
463) attempt to outline the history of scleromorphic flora may need qualification.
Central to their hypotheses was the idea that, following Specht (1979, 1981), the
scleromorphic flora originated by the early Palaeogene in the adaption to patches
of oligotrophic soils within forest vegetation. Secondly, they suggest that this
scleromorphic flora has a history as a unit through into its present prominence in
areas of Mediterranean climate and poor soils.

The evidence from southern Africa suggests otherwise. In the first place, it
appears that during the early Tertiary in the Arnot region members of the
Ericaceae, Restionaceae, Proteaceae and Thymelaeaceae (and perhaps Rubi-
aceae) grew on eutrophic soils within a lowland, probably extensive, forest,
growing under a warm dryish climate. Although local edaphic factors such as
forest disturbance (fire?), steep slopes or thin soil cover may have favoured their

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

growth, they must basically have been widespread within the forest, i.e. part of
the understorey. Secondly, this evidence taken together with (i) the dominance of
Restionaceae with graminoid apertures (unlike those recorded at Arnot and like
most extant southern African Restionaceae), (ii) the absence of Ericaceae in the
Knysna lignites (Thiergart et al. 1963) and (iii) the paucity of Restionaceae,
Ericaceae and Thymelaeaceae at Noordhoek (a site rimmed by mountains of
Table Mountain Sandstone, and therefore with very oligotrophic soils) in the
south-western Cape in the early Neogene (Coetzee 1978a, 1978b), suggests that
the origins of the present scleromorphic Capensis Flora are polyphyletic and that
the evolution of its components and history of its synthesis is complex.

Lastly, as suggested by Axelrod & Raven (1978) and Parrish et al. (1982),
relatively xeric vegetation may have a long history in the African subcontinent. If
this were so, comparative studies might show that relatively more of the older
scleromorphic taxa in the region were in fact truly sclerophyllous than is the case,
- for instance, in Australia.

21. The contribution of the Arnot evidence towards a preliminary outline of
the vegetation history of the African subcontinent during the late Cretaceous and
early Tertiary is discussed elsewhere (Coetzee et al. 1983). It suffices here to say
that at present the Arnot palynoflora records the first modern angiosperm flora
known in the African subcontinent after the extinction of the late Cretaceous,
archaic angiosperm flora in which Ephedripites, Fenestriorites, Cretacaeiporites,
Hexaporotricolpites and Proteacidites forms are prominent (Scholtz & Deacon
1982). The strong representation of forms such as Triorites operculatus,
T. sphericus and T. harrissii may relate to the often-recorded increase of triporate
forms with affinity to families such as the Betulaceae, Ulmaceae, Carpinaceae or
Casuarinaceae in the Palaeocene (Chourey 1974; Srivastava 1981). This phe-
nomenon has been taken, in conjunction with other evidence, to indicate a
cooling event in the early Palaeocene. Also, it has been suggested by Whitehead
(1971), amongst others, that the appearance of relatively small, triporate, psilate
angiosperm pollen indicating secondary adaptation to anemophily, coincides with
the appearance of the deciduous habit and seasonality of precipitation.

22. Lastly, the floral changes recorded within the Arnot sequence, as well as
the very different palynomorph assemblages known from the region and from
‘kimberlite’ pipe sequences from Botswana, raises a general point about the
possible differences that will be encountered in the study of vegetation change
based on ‘kimberlite’ pipe sequences, versus studies based on the more usual
depositional site sequences.

As already suggested, the crater lakes of ‘kimberlite’ volcanoes formed small
sedimentary traps positively unrelated to developed drainage patterns. It can be
assumed that their palynomorph assemblages were the result of wind and very
local water transport. In addition, the rate of their infilling is likely to have been
rapid. They will, therefore, reflect vegetation change to much finer scale than is
the case with larger epicontinental or deep-ocean sedimentary basins. This poses

PALYNOLOGY OF THE ARNOT PIPE 103

some problems for palynological work but, because of the abundance and
distribution of these sites, creates the potential for a detailed understanding of
vegetation associations, distribution and history in the time ranges concerned.

ACKNOWLEDGEMENTS

This research was funded as a project within a Council for Scientific and
Industrial Research—Cooperative Scientific Programmes (CSIR-—CSP) project,
the Fynbos Biome Project (Palaeoecology of the Fynbos Biome Subproject). The
research was originally started as part of input into the review paper “The
comparative evolution of Mediterranean-type ecosystems’ prepared by Professor
H. J. Deacon of the Department of Archaeology, University of Stellenbosch for
the CSIR-—CSP-funded conference on Mediterranean ecosystems (MEDCON
1980). The help and advice of Professor H. J. Deacon is gratefully acknowledged.

Mrs C. E. Stevens of the Department of Archaeology, University of
Stellenbosch, gracefully typed and retyped the manuscript, and painstaking care
was taken in producing the photomicrograph prints by Elsabé Pretorius. Many
thanks also to Mr A. P. and Mrs Christine Meyer, and to Esmien and Jannie
Louw of the farms Banke and Riembreek respectively, for their friendship and
hospitality.

Helpful criticism of the manuscript was received from Professor E. J. Moll,
Dr L. Scott and Dr E. M. Truswell (formerly Kemp). Dr M. Cluver of the South
African Museum kindly made the samples available for study. I am very grateful
to Miss E. Louw for considerable editorial assistance.

The University of Stellenbosch provided a generous grant towards part of the
publication costs of this work.

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Couper, R. A. 1953. Upper Mesozoic and Cainozoic spores and ‘pollen grains from New
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Couper, R. A. 1960. New Zealand Mesozoic and Cenozoic plant microfossils. Palaeont. Bull.,
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Daucuerty, L. H. 1941. The Upper Triassic flora of Arizona. Publs Carnegie Instn 526: 1-108.

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northeastern Brazil. Boln Asoc. latinoam. Paleobot. Palinol. 3: 14-20.

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Vict. 77: 1-148.

PALYNOLOGY OF THE ARNOT PIPE 105

DETIMANN, M. E. 1973. Angiospermous pollen from Albian to Turonian sediments of eastern
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Doy1eE, J. A. 1969. Cretaceous angiosperm pollen of the Atlantic coastal plain and its
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angiosperm pollen, with notes on the Cretaceous palynostratigraphy and palaeoenviron-
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Ersik, W. C. 1968. Palynology of a Paleocene Rockdale lignite, Milam Country, Texas. 1.
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ERDTMAN, G. 1938. Pollenanalys och pollenmorfologi. Nyr metodr och underskOningar. Svensk
bot. Tidskr. 32: 130-132.

ERDTMAN, G. 1952. Pollen morphology and plant taxonomy. Angiosperms. Stockholm: Almquist
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ERDTMAN, G. 1960. On three new genera from the lower Healdon Beds, Hampshire. Bot.
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Estes, R. 1977. Relationship of the South African fossil frog Eoxenopoides reuningi (Anura,
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FEerGuSON, I. K. 1977. Cornaceae. World Pollen & Spore Flora 6: 1-34.

Game_Erro, J. C. Morfologia del polen de Apterocladus lanceolata (Coniferae) de la Formacion
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GERMERAAD, J. H., Hoppinc, C. A. & MULLER, J. 1968. Palynology of Tertiary sediments of
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GoLbBLATT, P. 1978. An analysis of the flora of southern Africa: its characteristics, relationships
and origins. Ann. Mo. bot. Gdn 65: 369-436.

Harris, W. K. 1965. Basal Tertiary microfloras from the Princetown area, Victoria, Australia.
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Harris, W. K. 1974. Palynology of Paleocene sediments at Site 214, Ninety-east Ridge. Jnitial
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HASKELL, T. R. 1968. Saccate pollen grains from the lower Cretaceous of the great Artesian
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HEKEL, H. 1972. Pollen and spore assemblages from Queensland Tertiary sediments. Publs geol.
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HELGREN, D. M. & Burzer, K. W. 1977. Palaeosols of the southern Cape coast, South Africa:
implications for laterite definition, genesis and age. Geogrl Rev. 67: 430-445.

HERNGREEN, G. F. W. 1975. An upper Senonian pollen assemblage of borehole 3-PIA-10-AL,
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HERNGREEN, G. F. W. & CHLONOovA, A. F. 1981. Cretaceous microfloral provinces. Pollen
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HEusser, C. J. 1971. Pollens and spores of Chile. Tucson, Arizona: University of Arizona Press.

Hipeux, M. J. & Fercuson, I. K. 1976. The stereostructure of the exine and its evolutionary
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IprAHIM, H. D. 1933. Sporenformen des Aegirhorizonts der Ruhr-Reviers. Dissn tech. Hochsch.
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JANSONIUS, J. 1971. Emended diagnosis of Alisporites Daugherty, 1941. Pollen Spores 13:
349-357.

JANSONIUS, J. & Huis, L. V. 1976. Genera file of fossil spores and pollen. Calgary, Canada:
Department of Geology, University of Calgary.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

JARDINE, S. & MactorrE, L. 1963. Palynologie et stratigraphie du Crétacé des bassins du
Sénégal et de Cote d’Ivoire. Mém. Bur. Rech géol. miniér. 32: 187-245.

JARZEN, D. M. 1978. The terrestrial palynoflora from the Cretaceous—Tertiary transition,
Alabama, U.S.A. Pollen Spores 20: 535-553.

JARZEN, D. M. 1980. On the occurrence of Gunnera pollen in the fossil record. Biotropica 12:
117-123.

Jounson, L. A. S. & Briccs, B. G. 1981. Three old southern families—Myrtaceae, Proteaceae
and Restionaceae. Jn: KEast, A. ed. Ecological biogeography of Australia 1: 427-469. The
Hague: Junk.

Kepves, M. 1980. Morphological investigation of recent Palmae pollen grains. Acta bot. hung.
26: 339-373.

Kemp, E. M. 1978. Tertiary climatic evolution and vegetation history in the south-east Indian
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Kemp, E. M. 1981. Tertiary palaeogeography and the evolution of Australian climate. Jn:
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Kemp, E. M. & Harris, W. K. 1977. The palynology of early Tertiary sediments, Ninety-east
Ridge, Indian Ocean. Spec. Pap. Palaeont. 19: 1-69.

KIRCHHEIMER, F. 1934. On pollen from the upper Cretaceous dysodil of Banke, Namaqualand
(South Africa). Trans. R. Soc. S. Afr. 21: 41-50.

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Korova, I. Z. 1978. Spores and pollens from Cretaceous deposits of the eastern North Atlantic
Ocean, Deep Sea Drilling Project, Leg 41, Sites 367 and 370. Initial Rep. Deep Sea Drilling
Proj. 41: 841-881.

Kreme, G. O. W. 1965. Morphologic encyclopedia of palynology. Tucson: University of Arizona
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KRONER, A. 1973. Comments on ‘Is the African plate stationary?’ Nature, Lond. 243: 29-30.

Krutzscu, W. 1959. Mircopalaontologische (Sporen palaontologische) Untersuchungen in der
Braunkohle des Geiseltales. Geologie 8: (21/22): 1-425.

Lapp, P. G. 1977. Pollen morphology of some members of the Restionaceae and related families
with notes on the fossil record. Grana 16: 1-14.

Lams, H. H. 1972. Climate, present, past and future. 1. London: Methuen.

LEIDELMEYER, P. 1966. The Paleocene and lower Eocene pollen flora of Guyana. Leid. geol.
Meded. 38: 49-70.

Levyns, M. R. 1938. Some evidence bearing on the past history of the Cape Flora. Trans. R.
Soc. S. Afr. 26: 401-424.

Levyns, M. R. 1952. Clues to the past in the Cape Flora of today. S. Afr. J. Sci. 49: 155-164.

Levyns, M. R. 1964. Migrations and origins of the Cape Flora. Trans. R. Soc. S. Afr. 37:
85-107. :

Lewis, W. H. 1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia (Rubiaceae).
Am. J. Bot. 52: 257-284.

Lozano-GarciA, S. 1979. Atlas de polen de San Luis Potosi, Mexico. Pollen Spores 21:
287-336.

Masutrt, J. A. 1955. Erosion surfaces in Namaqualand and the ages of surface deposits in the
south-western Kalahari. Trans. geol. Soc. S. Afr. 58: 13-30.

MaaIL1, R. E. 1981. Bryophyta. Fascicle 1. Sphagnaceae—Grimmiaceae. Jn: LEISTNER, O. A.
ed. Flora of southern Africa. Pretoria: Botanical Research Institute, Department of
Agriculture & Fisheries.

Martin, A. R. H. 1959. South African palynological studies. 1. Statistical and morphological
variations in the South African species of Podocarpus. Grana 2: 40-53.

Martin, A. R. H. & Harris, W. K. 1974. Reappraisal of some palynomorphs of supposed
proteaceous affinity. The genus Proteacidites Cookson ex Couper. Grana 14: 108-113.
Martin, H. A. 1973. The palynology of some Tertiary and Pleistocene deposits, Lachlan River

Valley, New South Wales. Aust. J. Bot. (Suppl.) 6: 1-57.

~ Martin, H. A. 1978. Evolution of the Australian flora and vegetation through the Tertiary:
evidence from pollen. Alcheringa 2: 181-202.

Martin, H. A. 1981. The Tertiary flora. In: Keast, A. ed. Ecological biogeography of Australia
1: 391-406. The Hague: Junk.

PALYNOLOGY OF THE ARNOT PIPE 107

MatTHEws, J. M. 1966. Some descriptions of the pollen morphology of Epacris Forst., emend.
Cav. Pollen Spores 8: 461-478.

McIntyre, D. J. 1965. Some new pollen species from New Zealand Tertiary deposits. N. Z. J/
Bot. 3: 204-214.

McLacuian, I. R. & PieTerse. E. 1978. Preliminary palynological results: Site 361, Leg 40,
Deep Sea Drilling Project. Initial Rep. Deep Sea Drilling Proj. 4: 857-881.

Moorg, A. E. 1973. The olivine-melilitite association of Namaqualand. In: Extended abstracts of
the First International Kimberlite Conference, Cape Town, South Africa. Rondebosch:
University of Cape Town.

Moore, A. E. 1979. The geochemistry of the olivine melilitites and related rocks of Namaqua-
land/Bushmanland, South Africa. Unpublished Ph.D. thesis, University of Cape Town.
Morean, R. 1978. Albian to Senonian palynology of Site 364, Angola Basin. Initial Rep. Deep

Sea Drilling Proj. 40: 915-951.

MULLER, J. 1968. Palynology of the Pedawan and Plateau Sandstone formations (Cretaceous—
Eocene) in Sarawak, Malaysia. Micropalaeontologist 14: 1-37.

MULLER, J. 1980. Palynological evidence for Paleogene climatic changes. Mém. Mus. natn. Hist.
nat., Paris (B) 27: 211-218.

MULLER, J. 1981. Fossil pollen records of extant angiosperms. Bot. Rev. 47: 1-142.

MULLER-STOLL, W. R. & MADEL, E. 1962. Fossil woods of Monimiaceae and Euphorbiaceae
from the Upper Cretaceous Umzamba-beds of east Pondoland. Trans. geol. Soc. S. Afr. 65:
93-104.

Muri1o, M. T. & Biess, M. J. M. 1974. Spores of Recent Columbian Pteridophyta. 1. Trilete
spores. Rev. Palaeobot. Palynol. 18: 223-269.

Muritto, M. T. & Bress, M. J. M. 1978. Spores of Recent Columbian Pteridophyta. 2.
Monolete spores. Rev. Palaeobot. Palynol. 25: 319-365.

Nicuois, D. J., Ames, H. T. & TRAveRSE, A. 1973. On Arecipites Wodehouse, Monocolpo-
pollenites Thomson & Pflug, and the species Monocolpopollenites tranquilus. Taxon 22:
241-256.

Nitsson, S., PRAGLOWSKI, J. & NiLsson L. 1977. Airborne pollen grains and spores in northern
Europe. Stockholm: Bokforlaget Natur och Kultur.

ParrisH, J. T. & Curtis, R. L. 1982. Atmospheric circulation, upwelling and organic-
rich rocks in the Mesozoic and Cenozoic eras. Palaeogeogr. Palaeoclimat. Palaeoecol. 41:
31-66.

ParrisH, J. T., ZIEGLER, A. M. & ScoresE, C. R. 1982. Rainfall patterns and the distribution of
coals and evaporites in the Mesozoic and Cenozoic. Palaeogeogr. Palaeoclimat. Palaeoecol.
41: 67-101.

PocKNALL, D. T. Pollen morphology of the New Zealand species of Dacrydium Solander,
Podocarpus L’Heritier and Dacrycarpus Endlicher (Podocarpaceae). N. Z. Jl Bot. 19:
67-95.

Poo.e, M. M. 1981. Pollen diversity in Zimmermania (Euphorbiaceae). Kew Bull. 36: 129-138.

PoToni£, R. 1956. Synopsis der Gattungen der Sporae dispersae, Pt. 1. Beih. geol. Jb. 23: 1-103.

Punt, W. 1962. Pollen morphology of the Euphorbiaceae with special reference to taxonomy.
Wentia 7: 1-116.

ReitsMA, T. J. 1970. Pollen morphology of the Alangiaceae. Rev. Palaeobot. Palynol. 10:
249-332.

RENNIE, J. V. L. 1931. Note on fossil leaves from the Banke clays. Trans. R. Soc. S. Afr. 19:
251-253.

REUNING, E. 1931. A contribution to the geology and palaenotology of the western edge of the
Bushmanland plateau. Trans. R. Soc. S. Afr. 19: 215-232.

REUNING, E. 1934. The composition of the deeper sediments of the pipe at Banke,
Namaqualand, and their relation to kimberlite. Trans. R. Soc. S. Afr. 21: 33-39.

ReyreE, Y. 1973. Palynologie du Mesozoique saharien. Mém. Mus. natn. Hist. nat., Paris (C) 27:
1-284.

Rocers, A. W. 1911. Geological survey of parts of Van Rhynsdorp and Namaqualand Divisions.
Rep. geol. Commn Cape Good Hope 16: 9-84.

SAAD, S. I. & GHAZALY, G. 1976. Palynological studies in Nubia Sandstone from Kharg Qasis.
Pollen Spores 18: 407-470.

Sau, S. C. D. 1967. Palynology of an upper Neogene profile from Rusizi Valley (Burundi). Annis
Musée r. Afr. cent. Ser. 8vo (Sci. géol.) 57: 1-173.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

SALARD-CHEBOLDAEFF, M. 1978. Sur la Palynoflore Maestrichtienne et Tertiaire du bassin
sédimentaire littoral du Cameroun. Pollen Spores 20: 213-260.

SALARD-CHEBOLDAEFF, M. 1979. Palynologie Maestrichtienne et Tertiaire du Cameroun. Etude
qualitative et repartition verticale des principales espéces. Rev. Palaeobot. Palynol. 28:
365-388.

SALARD-CHEBOLDAEFF, M. 1981. Palynologie Maestrichtienne et Tertiaire du Cameroun.
Résultats botaniques. Rev. Palaeobot. Palynol. 32: 401-439.

SAXENA, R. K. 1982. Taxonomic study of the polycolpate pollen grains from the Indian Tertiary
sediments. Rev. Palaeobot. Palynol. 37: 283-316.

ScHoLttz, A. & Deacon, H. J. 1982. Report on fossil pollens from sediments associated with a
group of kimberlite pipes in Botswana. Internal Report, Department of Archaeology,
University of Stellenbosch, to a diamond mining company.

Scott, L. 1976. Palynology of Lower Cretaceous deposits from the Algoa Basin (Republic of
South Africa). Pollen Spores 18: 563-609.

SENESSE, S. 1980. Palynologia Madagassica et Mascarenica. Fam. 98 bis: Caesalpinaceae. Pollen
Spores 22: 355-423.

Sowumnl, M. A. 1972. Pollen morphology of the Palmae and its bearing on taxonomy. Rev.
Palaeobot. Palynol. 13: 1-80.

SpecHuT, R. L. 1979. Heathlands and related shrublands of the world. In: Specnt, R. L. ed.

Ecosystems of the world. Vol. 9. Heathlands and related shrublands: 1-18. Amsterdam:
Elsevier.

SpEcHT, R. L. 1981. Evolution of the Australian flora: some generalisations. Jn: KEAsT, A. ed.
Ecological biogeography of Australia 1: 783-806. The Hague: Junk.

SRIVASTAVA, S. K. 1969. Assorted angiosperm pollen from the Edmonton Formation
(Maestrichtian), Alberta, Canada. Can. J. Bot. 47: 975-989.

SRIVASTAVA, S. K. 1971. Monolete spores from the Edmonton Formation (Maastrichtian),
Alberta (Canada). Rev. Palaeobot. Palynol. 11: 251-265.

SRIVASTAVA, S. K. 1981. Evolution of upper Cretaceous phyto-geoprovinces and their pollen
flora. Rev. Palaeobot. Palynol. 35: 155-173.

Stover, L. E. & Evans, P. R. 1973. Upper Cretaceous—Eocene spore-pollen zonation offshore
Gippsland Basin, Australia. Spec. Publs geol. Soc. Aust. 4: 55-72.

STovER, L. E. & PARTRIDGE, A. D. 1973. Tertiary and Late Cretaceous spores and pollen from
the Gippsland Basin, southeastern Australia. Proc. R. Soc. Vict. 85: 237-286.

TAKHTAJAN, A. 1969. Flowering plants. Origin and dispersal. Edinburgh: Oliver & Boyd.

TANKARD, A. J. & RoGers, J. 1978. Late Cenozoic palaeoenvironments on the west coast of
southern Africa. J. Biogeogr. 5: 319-337.

Tay Lor, H. C. 1978. Capensis. In: WERGER, M. J. A. ed. Biogeography and ecology of southern
Africa, 1: 171-229. The Hague: Junk.

THIERGART, F., FRANTz, U. & Rauxopr, K. 1963. Palynologische Untersuchungen von
Tertiarkohlen und einer oberflachen Probe nahe Knysna, Siid-Afrika. Advg Front. Pl. Sci.
4: 151-178.

THomson, P. W. & Priuc, M. 1953. Pollen und Spores des mitteleuropaischen Tertiars.
Palaeontographica (B) 94: 1-138.

TRUSWELL, E. M. & Harris, W. K. 1982. The Cainozoic palaeobotanical record in arid
Australia: fossil evidence for the origins of an arid-adapted flora. Jn: PARKER, W. R. &
GREENSLADE, P. J. M. eds. Evolution of the flora and fauna of arid Australia: 67-76.
Adelaide: Peacock Publications.

VAN DER HAMMEN, T. 1954. El desarrolo de la flora Colombiana en los periodos geologicos, 1.
Maastrichtiano hasta Terciario mas inferior. Boln geol., Bogota 4: 63-101.

VAN HOEKEN-KLINKENBERG, P. M. J. 1964. A palynological investigation of some Upper
Cretaceous sediments in Nigeria. Pollen Spores 6: 34-48.

VAN HOEKEN-KLINKENBERG, P. M. J. 1966. Maastrichtian, Paleocene and Eocene pollen and
spores from Nigeria. Leid. geol. Meded. 38: 34-48.

Wess. L. J. & Tracey. J. G. 1981. Australian rainforests: patterns and change. Jn: KEastT, A.
ed. Ecological biogeography of Australia 1: 605-691. The Hague: Junk.

WELMAN, W. G. 1970. The South African fern spores. In: VAN ZINDEREN BAKKER, E. M. ed.
South African pollen grains and spores. Part 6. Cape Town: Balkema.

WHITEHEAD, D. R. 1971. Wind pollination in the angiosperms: evolutionary and environmental
considerations. Evolution 23: 28-35.

PALYNOLOGY OF THE ARNOT PIPE 109

WILLIis, J. C. 1966. A dictionary of the flowering plants and ferns (7th ed.). Cambridge:
Cambridge University Press.

WILSon, L. R. & WEBSTER, R. M. 1946. Plant microfossils from a Fort Union coal of Montana.
Am. J. Bot. 33: 271-278.

WoDEHOUSE, R. P. 1933. Tertiary pollen. II. Pollens of the Green River oil shales. Bull. Torrey
bot. Club 60: 479-524.

WopPENER, H., CALLEN, R. & Harris, W. K. 1974. The lower Tertiary Eyre Formation of the
southwestern Great Artesian Basin. J. geol. Soc. Aust. 21: 17-51.

7 & “2
n
: = .
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= = t 7
aa i 1—& a
= 8 x ri Se =
r F a -
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ao
ae
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)
ae i a -
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} : ;
a = SSS Se —_—_—SSSSES__S__Co ee ees = = = SSS = = ———— —— ——— —

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syN. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters
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A. SCHOLTZ

THE PALYNOLOGY OF THE
UPPER LACUSTRINE SEDIMENTS
OF THE ARNOT PIPE,

BANKE, NAMAQUALAND

VOLUME 95 PART 2 APRIL ISSN 0303-2515

CAPE TOWN

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, p. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHer, P.-H., DuvAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, cee s masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): Sil.

THIELE, J. 1910. Mollusca: B. Pelyyneaahont Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
April 1985 April
Part 2Z Deel

THE FAUNAL DEPOSITS OF A
LATE PLEISTOCENE RAISED BEACH
AT MILNERTON, CAPE PROVINCE,
SOUTH AFRICA
By
BRIAN KENSLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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word uitgegee in dele op ongereelde tye na gelang van die
beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

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GCL, iso), TID) & O02, m), IOS),
IN(_2, 5, 7, tapi), 142), 15425), 2400), 27: 3103), 326) 33, soe naaen

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ISBN 0 86813 065 6

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE FAUNAL DEPOSITS OF A LATE PLEISTOCENE
RAISED BEACH AT MILNERTON, CAPE PROVINCE, SOUTH AFRICA

By
BRIAN KENSLEY

Smithsonian Institution, Washington, D.C.

(With 2 figures and 3 tables)

[MS accepted 21 March 1984]

ABSTRACT

The faunal content of a Late Pleistocene raised beach exposed on the north shore of Table
Bay is examined. The deposit has been correlated with the Velddrif Shelly Sand Member of the
Bredasdorp Formation. The deposit contained mainly molluscan shells (78 species), with
occasional crustacean, echinoderm, and elasmobranch fish remains. The molluscs represent
rocky-shore, sandy-shore, and calm-water and/or estuarine species. It is hypothesized that this
mixed assemblage is due to a kill-off (perhaps because of a cut-off from the sea and rising salinity
and temperatures) in a nearby lagoonal area (the Rietvlei Basin), with the dead shells eventually
being washed out to sea, and then thrown up at the top of the beach, along with the remains of
sandy-beach and rocky-shore forms. The deposit contains two extinct species, Nuculana
bicuspidata and Crepidula capensis praerugulosa, as well as 12 species now confined to the
warmer waters of the east coast.

CONTENTS

PAGE
MiNtHO CU GtOMMPE, oe ye he race ese onde a ee ents Be tat
JAVA OG IS “ej Som RR ew eI rit ge Se eam 2
DEScHIpPUOMmOthe deposit. . 52... 6. --s ane sehen see eee 112
ENGSULIG HPP R ert geiite: fe De a hE Ue ee See? ees Oe 114
DISCUSSION forte te tre ee he Aan TAS ete el eae nae ae 121
PRCKMOWLEGE CIMEM(Sie is ses ee ei liga bk aoe eat eee 122
IGIONCT COSHe Pi ee ey ert, A ee, Sere re ee 12

INTRODUCTION

In June 1974, during the heavy winter weather experienced in Table Bay, a
north-west storm coincided with a spring-tide. The resultant exceptionally high
and powerful wave action eroded a section of the beach and fringing sand-dunes
just below the Milnerton lighthouse, and exposed a sedimentary deposit
dominated by molluscan shells. In May 1983, a short stretch of an old beach-line,
about 0,5 km south of the lighthouse on the Cape Town side of the Milnerton
Lagoon mouth, was exposed. Superficial inspection of the deposits revealed
several features that pointed to a Pleistocene age. These included a very obvious
concentration of molluscan shells, the brown colour of what was obviously the
common black mussel Choromytilus meridionalis, and the presence of species
that do not now occur alive in Table Bay or on the west coast of southern Africa.
The object of this paper is to place the deposits and their probable age on record,
and to speculate on their history.

JUL

Ann. S. Afr. Mus. 95 (2), 1985: 111-122, 2 figs, 3 tables.

1D ANNALS OF THE SOUTH AFRICAN MUSEUM

METHODS

To determine species composition, selective manual collecting was done
along the deposit, and a faunal list drawn up.

In an attempt to gain a rough idea of the quantitative composition, a cubic
metre of deposit was collected, washed in water to separate the fossils, and
species and specimens sorted, identified and counted.

From molluscan shells supplied to Teledyne Isotopes of New Jersey, a
radiocarbon date was obtained (sample number I—8372).

DESCRIPTION OF THE DEPOSIT

The major exposed deposit is situated about 100 m to the north of Milnerton
lighthouse on the shore of Table Bay (33°53’S 18°27’E) (Figs 1, 2A—B). At the
Low Water of Springs level the beach was scoured away to expose a bed of
-ferricrete that showed a characteristic nodular and cellular structure (Fig. 2C).
Into the irregularities of this ferricrete, shells and coarse sediment had become
cemented. Where the shells actually touched the ferricrete, they were stained a
rusty brown. It is possible that this ferricrete layer is homologous with the ‘iron-
stained gravelly sands’ described by Tankard (1975a: 261) from a late Tertiary
deposit at Ysterplaat about 4 km away. In places in the lower part of the deposit,
patches of black peat-like material were exposed. Shells were not present in this
peat (Fig. 2E).

The whole area of the beach between Low Water of Springs and the
exceptionally high High Water of Springs revealed shell remains (Fig. 2F). In
places, the consolidating sediment seemed harder or more firmly cemented than
in others, and here lumps of the deposit that had eroded more slowly than the
softer sediments protruded above the more level ‘beach’ surface. At the top of the
beach, which normally is a gentle sand slope running into low sand-dunes, the sea
had cut a cliff into the bases of these dunes, exposing a vertical face in the deposit
of about 1 m in thickness. In places, this face of the deposit was interrupted by
gulleys of black non-fossiliferous sand (Fig. 2D). Both shell deposit and black
sand were overlain by modern, white, calcareous, littoral sand. Three weeks after
the sudden exposure of this deposit, all sign of it had vanished, having been
covered by white sand moved in by sea and wind.

The length of the major deposit exposed along the beach was 64 m. The
horizontal width of the beach from LWS to the top of the sand-dune cliff was
13 m. The vertical distance from LWS to the top of the deposit was 2,5 m.

The deposit consisted of coarse sand grains and shell debris, with occasional
angular rounded pebbles, and a few scattered pieces of calcrete. There was some
bedding, with especially the bivalve shells oriented horizontally, but this was not
everywhere apparent. |

The 1983 beach-line exposure south of the lagoon mouth consisted of a
40-50 cm-thick layer of calcrete containing sparsely scattered shells showing no
obvious bedding. Thin lenses of shells about 20 cm below the limestone could

A LATE PLEISTOCENE RAISED BEACH 113

e VELDDRIF

SALDANHA Berg Rive*

Saldanha Bay

ATLANTIC OCEAN

Rietvlei lj

major site

Milnerton
Lighthouse

Fig. 1. Map showing location of Milnerton beach deposit.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

occasionally be seen. While close to the mouth of the Milnerton Lagoon, this
deposit cannot be confused with the late Tertiary marine sediments referred to by
Tankard (1975a: 262) as ‘submerged deposits just offshore from Milnerton which
are below normal wave erosion base .

RESULTS

AGE OF THE DEPOSIT

A radiocarbon date of 33 750 + 1 780 years Bp was obtained, but this may be
a minimum age. The deposits have been correlated with the Velddrif Shelly Sand
Member of the Bredasdorp Formation (Tankard 1976) by Rogers (1982).

FAUNAL ANALYSIS

Table 1 gives the list of 78 species of molluscs, five other invertebrates, and
two vertebrates, found both in the cubic metre of deposit and in material hand-
collected at random. Records of the Quaternary occurrences of the species as well
as the present distribution are given, along with a rough indication of the
ecological habitat of each species.

The habitat types of the species may be sorted roughly into rock-dwelling
forms, sand or mud-dwellers, and estuarine and/or calm-water forms. (This latter
group is not more stringently divided for reasons both of definition, and because
little is known of the biology of several of the living forms.) From Table 1 it can be
seen that of these habitat-types, the greatest number of species as well as
specimens belong to the rock-dwelling group. The majority of these are forms
that occur to varying degrees of abundance in the intertidal zone. Seven species of
the estuarine and/or calm-water group, representing 7,6 per cent of the total
sample, were present. The most abundant species was an extinct Crepidula,
closely followed by an extant species of the same genus (see Table 2). The next
ten most abundant species are all living forms found on the west coast. Five
species are typical rock-dwelling forms, five species are sand or mud-dwellers,
and amongst these latter are forms that occur in sandy habitats exposed to strong
wave action, e.g. Bullia digitalis, as well as forms that occur in either sublittoral or
calm water, e.g. Bullia laevissima, Nassarius speciosus.

Species that do not occur living at the present on the west coast are also
represented in the deposit. This gives a list of 12 species, all typical inhabitants of
the warmer waters of the south-east and east coasts (see Table 3). Of these 12
species, six have been recorded from the Pleistocene deposits of the west coast,
mainly from the Elands Bay—Velddrif—Saldanha Bay area (see Tankard 1975);
Schalke 1973; Visser & Schoch 1973; Barnard 1962).

Given the probable Eemian Interglacial age for the deposit, it would not be
unreasonable to expect (in the light of Pleistocene molluscan extinctions) a few
extinct forms in the present assemblage. One extinct species is present, plus one
species no longer occurring live in southern Africa. Nuculana bicuspidata, a

A LATE PLEISTOCENE RAISED BEACH 115

Fig.2. A. Milnerton beach, looking north, showing shell deposit and overlying sand dunes at right. B. Milnerton beach,
looking south towards Cape Town, showing shell deposit. C. Ferricrete exposed at lower level of beach. D. Non-
fossiliferous dark sand below white dune sand. ___E. Peat-like material in shell deposit. F. Close-up of shell deposit.

ANNALS OF THE SOUTH AFRICAN MUSEUM

116

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E—estuarine; M—mud dweller; R—rocky-shore dweller; S—sand dweller: W—weed-bed dweller: +—extinct speci
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Species

Mo tuscA, BIVALVIA
Aulacomya ater (Molina)

Barnea truncata Say
Choromytilus meridionalis (Krauss)
Donax serra (Chemnitz)

Dosinia lupinus Linnaeus
Loripes liratula (Sowerby)

Lutraria lutraria (Linnaeus)
Mactra glabrata Linnaeus

Melliteryx mactroides (Hanley)
Nuculana biscuspidata (Gould)
Ostrea algoensis Sowerby
Parvicardium turtoni (Sowerby)
Petricola bicolor Sowerby
Psammotellina capensis Sowerby

Scissodesma spengleri (Linnaeus)
Solen capensis Fischer

Tellimya trigona Barnard
Tellina trilatera Gmelin

Theora ovalis Smith
Tivela compressa (Sowerby)

Venerupis corrugata (Gmelin)

MOoLLuscaA, GASTROPODA

Afrocominella capensis ( Dunker)
Amblychilepas scutellum ( Gmelin)

Argobuccinum pustulosum (Lightfoot)

Assiminea globulus Connolly
Bullia annulata (Lamarck)
Bullia digitalis Meuschen

Bullia laevissima (Gmelin)

Burnupena cincta (Roding)
Burnupena lagenaria (Lamarck)

Burnupena papyracea (Bruguiere)

Calyptraea chinensis (Linnaeus)

Clionella confusa (Smith)

Cinysca granulosa (Krauss)

Conus mozambiecus Hwass

Crepidula capensis praerugulosa Kilburn
& Tankard

Crepidula porcellana Lamarck

Crepidula rugulosa Dunker
Cymatium cutaceum africanum (Adams)

Cythara amplexa (Gould),
Epitonium kraussi (Nyst)
Fissurella mutabilis Sowerby

Gibbula capensis (Gmelin)
Gibbula cicer (Menke)

Helcion dunkeri (Krauss)
Lippistes cornu (Gmelin)
Littorina knysnaensis (Philippi)
Marginella rosea Lamarck

TABLE 1

Faunal list, Milnerton Late Pleistocene raised beach.

Quaternary records

Liideritz, Orange River, Velddrif, Saldanha,
Rietvlei

Table Bay
Orange River, Sedgefield, Durban

Liideritz, Orange River, Velddrif, Saldanha,
Langebaanweg, Sedgefield

Cape Cross, Saldanha, Velddrif, Bredas-
dorp, Sedgefield, Port Elizabeth

Saldanha, Little Brak River, Sedgefield,
Knysna

Liideritz, Saldanha, Velddrif, Sedgefield

Saldanha, Little Brak River, Sedgefield, Port
Elizabeth, Durban

Cape Cross, Velddrif

Elands Bay, Velddrif, Knysna

Sedgefield, Knysna, Port Elizabeth, Durban

Elands Bay, Velddrif, Saldanha, Rietvlei,
Sedgefield

Velddrif, Saldanha, Port Elizabeth

Lideritz, Elands Bay, Velddrif, Saldanha,
Rietvlei, Sedgefield, Port Elizabeth

Elands Bay, Velddrif, Saldanha, Rietvlei

Orange River, Velddrif, Saldanha, Little
Brak River, Knysna

Velddrif, Saldanha, Bredasdorp, Little Brak
River

Orange River, Elands Bay, Velddrif, Sal-
danha, Little Brak River, Sedgefield

Velddrif, Saldanha, Little Brak River,
Sedgefield, Inhambane

Liideritz, Orange River Mouth, Velddrif,
Saldanha

Saldanha, Port Elizabeth

Liideritz, Orange River, Velddrif, Saldanha,
Port Elizabeth

Cape Cross, Liideritz, Velddrif, Saldanha,
Sedgefield, Port Elizabeth

Elands Bay, Velddrif, Saldanha

Liideritz, Orange River, Saldanha, Port
Elizabeth

Velddrif, Saldanha

Velddrif, Saldanha, Port Elizabeth

Orange River, Saldanha
Elands Bay, Velddrif, Saldanha

Liideritz, Orange River, Velddrif, Saldanha,
Knysna, Port Elizabeth

Liideritz

Velddrif, Saldanha, Little Brak River,
Sedgefield, Knysna, Port Elizabeth,
Durban

Velddrif, Saldanha

Saldanha, Rietvlei, Little Brak River, Port
Elizabeth

Saldanha

Bredasdorp

Elands Bay, Saldanha, Knysna

Living distribution

Habitat
Namibia to Natal R
Senegal to Angola R
Namibia to Natal Rins
Namibia to Port Alfred Ss
Walvis Bay to East London S&M
Still Bay to Port Alfred S
Luderitz to Port Alfred S
Saldanha to Natal S
False Bay to Port Alfred S
Mauritania to Angola S
False Bay to East London R
False Bay to Natal S
Namibia to Zululand R
False Bay to Port Alfred E
False Bay to Port Alfred S
Olifants River to East London M,E
Liideritz to False Bay S
Saldanha to Port Alfred S
Saldanha to Port Alfred S
False Bay to Natal S
West Africa to Natal S.R
Namibia to Cape Agulhas R
Angola to Natal R
Namibia to East London R
Olifants River to Keiskamma River M,E
Saldanha to Mozambique S
Namibia to Transkei S
Namibia to Transkei S,M
Angola to Transkei R
Namibia to Natal R
Namibia to Walker Bay R
Saldanha to East London R
Mossel Bay to Port Alfred R
Namibia to Transkei R
Namibia to East London R
+ 2R
North-west Africa to Natal R
Lamberts Bay to Still Bay R
Namibia to Mozambique R
Liideritz to East London R
Namibia to Natal R
Liideritz to Natal R
Saldanha to Agulhas R
Namibia to Transkei R
Namibia to Natal R
Table Bay, East London ?
Namibia to Natal R
R,inS

Saldanha to Agulhas

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“TOATY, YI 9p] “eyUuepyes “JUpplsA

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ueqing ‘yleqezyq 110g
‘euskuy ‘playaspas ‘1oary yesg
apa] ‘diopsepoaig ‘eyuepyes “JUpploA.

yoqezi[q 140g

sp1ode1 Areuio}end

(ssnery) sisuadvo vyjawojodg

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(uOJUY) DIDSa1IDA ajajskxE
(zyUWIAYD) vu1431] ajaiskxE

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(jorewley]) vsowonbs pvjyjaonn

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UOJUY JIA] VIIDN
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(A9yUNC) SNUDISSNDAY SNIAVSSDN
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saroads

119

A LATE PLEISTOCENE RAISED BEACH

OE oe pe ee ee ee ee eee

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wan

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eyuepyes “YIO[ION Hod

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euskuyy

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VLVIHONVUAOWSVIA “SAOSIG

(aysaq) snsojnsun snuiyradvg “Jo
VGIONIHOYW ‘VLVNAaAGONIHOY

(ueepy op) vinjound sadyvacC
ZuIQqaIs Issnb4y VSspUdI]VD
vdaodvoaqd ‘VaAOVISNAD

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VUNANIHANY ‘VOSATIOW

(ssnery) s7suadpo pulipajoA
‘ds snjauia,

YOIeUIeT] DAafIUIADI VIJANAAN TL
SnoevuUry] SNINDUWADS OGAN
(1oyUNd) vUIIaU v1 0914]
(1oyund) sisuadns vor

‘ds piupuoydig

AqdIaMog sisuadvd vOSSIY
(q1gINsNIg) VjINIvIUNA DSNIAY
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(yy) 1pasfjo vuonydosopnasd

Species

Marginella sp.
Nassarius capensis (Dunker)
Nassarius kraussianus (Dunker)

Nassarius speciosus (Adams)
Natica saldontiana Bartsch
Natica tecta Anton

Nucella cingulata (Linnaeus)
Nucella dubia (Krauss)

Nucella squamosa (Lamarck)

Ocenebra scrobiculata (Philippi)
Oxystele tigrina (Chemnitz)
Oxystele variegata (Anton)
Patella argenvillei Krauss

Patella barbara Linnaeus
Patella granatina Linnaeus
Patella miniata Born

Protomella capensis (Krauss)

Pseudoraphitoma alfredi (Smith)
Pteropurpura uncinaria ( Lamarck)
Retusa truncatula (Bruguiére)
Rissoa capensis Sowerby
Siphonaria sp.

Tricolia capensis (Dunker)
Tricolia neritina (Dunker)

Turbo sarmaticus Linnaeus
Turritella carinifera Lamarck

Vermetus sp.

Volvarina capensis (Krauss)

Mo tiusca, AMPHINEURA

2 species represented by loose valves
CRUSTACEA, CIRRIPEDIA

Balanus maxillaris Gronovius
Balanus sp.

Crustacea, DECAPODA
Callianassa kraussi Stebbing
Ovalipes punctata (de Haan)
EcHINODERMATA, ECHINOIDA

ef. Parechinus angulosus (Leske)
Pisces, ELASMOBRANCHIATA
Myliobatis sp.

Odontaspis acutissima Agassiz

Quaternary records

Living distribution

Port Elizabeth

Velddrif, Saldanha, Bredasdorp, Little
Brak River, Sedgefield, Knysna,
Port Elizabeth, Durban

Velddrif, Saldanha
Velddrif, Saldanha

Velddrif, Saldanha, Little Brak River,
Sedgefield, Knysna, Port Elizabeth

Liideritz, Orange River, Velddrif, Saldanha

Orange River, Saldanha, Velddrif, Little
Brak River, Sedgefield, Port Elizabeth

Liideritz, Orange River, Velddrif, Saldanha,
Rietvlei

Elands Bay, Saldanha, Port Elizabeth
Elands Bay, Saldanha, Port Elizabeth

Liideritz, Orange River, Velddrif, Saldanha,
Port Elizabeth

Liideritz, Orange River, Velddrif, Saldanha
Liideritz, Orange River, Velddrif, Saldanha
Orange River, Velddrif, Saldanha

Velddrif, Saldanha, Rietvlei, Little Brak
River, Sedgefield, Knysna

Knysna
Elands Bay, Velddrif, Saldanha

Rietvlei

Port Nolloth, Saldanha
Orange River, Saldanha, Little Brak River,
Sedgefield, Knysna

Velddrif, Saldanha, Rietvlei

Velddrif

Saldanha, Milnerton

Table Bay to Transkei
Namaqualand to Mozambique

Orange River to Transkei
Saldanha to Agulhas
Namibia to East London

Namibia to False Bay
Namibia to Natal

Namibia to Transkei

Lideritz-to Transkei
Saldanha to Transkei
Angola to Natal

Namibia to Transkei

Namibia to Zululand
Namibia to Walker Bay
Namibia to Natal

Lamberts Bay to East London

Table Bay to Durban

False Bay to Natal

Agulhas to Port Alfred

Still Bay, Port Alfred
Namibia to Mozambique
Namibia to Port Elizabeth
Table Bay to Transkei

Port Nolloth to Mozambique

Liideritz to Agulhas

Lamberts Bay to Port Elizabeth

Olifants River to Natal
Namibia to Natal

Namibia to Zululand

os

Habitat

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20 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Ten major components of the Milnerton beach deposit,
as percentages of the total number of specimens.

Species Percentage
Crepidula capensis praerugulosa YD
Crepidula porcellana oll
Bullia laevissima 8,4
Bullia digitalis D5
Choromytilus meridionalis Sell
Tricolia capensis 4,8
Nassarius speciosus 39)
Nucella squamosa 3,8
Rissoa capensis 2,9
Venerupis corrugata 2,8

palaeotaxodont bivalve, has been reported as a Pleistocene fossil from Velddrif,
Cape Province, and Cape Cross, South West Africa (Namibia), but living from
Angola to Mauritania and the Cape Verde Islands (Tankard 19755). The animal
lives in sand or mud, probably infratidally, as it has been recorded from the same
sediments as Panopea glycymeris (Kensley 1974). Crepidula capensis praerugu-
losa Kilburn & Tankard, 1975, the most abundant species from the deposit, is also
known from an older Late Pleistocene site near Langebaan (9,5 ma.s.1.). Kilburn
& Tankard (1975) note that C. c. praerugulosa has a regularly curved ventral
margin, and speculate that the animals were often attached to mytilid mussels,

TABLE 3

Typical south-east and east-coast species occurring in the Milnerton deposit, with Pleistocene

Species

Clionella confusa
Loripes liratula

Nassarius capensis
Ostrea algoensis
Parvicardium turtoni
Psammotellina capensis
Pseudoraphitoma alfredi
Pteropurpura uncinaria
Retusa truncatula

Rissoa capensis
Scissodesma spengleri
Tivela compressa

records where known.

Pleistocene records

Elands Bay, Velddrif, Saldanha, Rietvlei, Little Brak River,
Keurbooms, Knysna

Algoa Bay

Elands Bay, Velddrif, Knysna

Sedgefield, Knysna, Algoa Bay, Durban

Elands Bay, Velddrif, Saldanha, Rietvlei, Sedgefield

Knysna

Elands Bay, Velddrif, Saldanha

Velddrif, Saldanha, Port Elizabeth

Velddrif, Saldanha, Bredasdorp, Little Brak River

A LATE PLEISTOCENE RAISED BEACH 121

also an occasional habitat for C. porcellana. As this latter species was the second
most abundant in the Milnerton deposit, it seems unlikely that they shared a
common habitat. Quite possibly, C. porcellana was more frequently attached to
large gastropods or to rocky substrates and, with the extinction of C. c. prae-
rugulosa, came to live on mussels as well. A third extinct form is the elasmo-
branch fish Odontaspis acutissima represented by a single unworn tooth. This
species is well recorded from the Pleistocene (Darteville & Casier 1943), but little
significance can be attached to its presence in the Milnerton deposit, as this single
shark tooth could be a reworked entity of earlier age.

Several features concerning the actual state of the shells in the deposit give
rise to comment, e.g. the concentration of shells, the condition of the shells, and
the abundance of shells less than 10 cm in length. The cubic metre of deposit
yielded 4 908 recognizable specimens comprising 82 species. While worn and
broken specimens were present, the majority were unworn and in perfect
condition, including many shells of small species or delicate juvenile shells.

DISCUSSION

At the time of deposition of the Milnerton accumulation, there must have
been an estuarine and/or calm-water environment nearby, supporting a more
diverse molluscan population than does the present-day Milnerton lagoon-—
estuary complex (cf. Millard & Scott 1954). This was probably the Rietvlei Basin,
which during the Pleniglacial opened north of both the present mouth and the
beach deposit under discussion (Schalke 1973). (Although from the faunal list the
Milnerton deposit appears richer and more diverse than Schalke’s Rietvlei fauna,
his sampling by means of bore-holes was obviously limited, and the two deposits
cannot usefully be compared.)

Also in the vicinity there must have been an extensive intertidal rocky-shore
area to support the relatively high numbers of rocky-shore forms, as well as a
sandy infra- and intertidal area.

The 12 species from the deposit now found living only on the east coast seem
to indicate that the sea temperature at the time of deposition was somewhat
higher than is presently experienced in Table Bay. Nine of the 12 species range
from False Bay eastwards, indicating a minimum sea temperature of 14 °C. The
present temperature regime in Table Bay has minima of 9-10°C. That
Pleistocene sea temperatures were at times higher than those of the present on
the west coast has already been well documented (Tankard 1975b). As a typical
west-coast fauna was already established at the time of the Milnerton deposition,
it seems probable that the 12 east-coast species listed in Table 3 were either
ecologically more tolerant than many of the west-African and east-coast forms,
which had already died out, or they represented the tail-end of populations dying
out on the southern west coast. ©

Schalke (1973) postulated a fluctuating rainfall pattern at about the time of
the deposition of the shells under discussion. A series of events that could lead to

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

the inclusion of numerous estuarine forms in the beach deposit would involve a
dry period resulting in a drying up and an increased temperature and salinity
regime of the estuary and/or calm-water area (assuming a temporary cut-off from
the sea, as during neap tides), which in turn would lead to large-scale mortality of
many of the physiologically less tolerant molluscs. With the advent of strong
rains, these dead shells would then be washed out to sea, to be deposited along
with the purely marine forms. This series of events has been witnessed on the
South West African coast (Kensley 1978), and could account to some degree for
the mixed nature of the Milnerton faunal assemblage.

ACKNOWLEDGEMENTS

I am grateful to Dr Q. B. Hendey, South African Museum, and
Dr J. Rogers, C.S.I.R. Marine Geology Unit, University of Cape Town, for their
advice and comments. My thanks are due to the Director and Trustees of the
South African Museum for permission to work on the material described during
my 1983 visit.

REFERENCES

BARNARD, K. H. 1962. Revised list of South African Late Tertiary and Pleistocene marine
Mollusca. Trans. R. Soc. S. Afr. 36: 179-196.

DARTEVILLE, C. & CASIER, E. 1943. Les poissons fossiles du Bas-Congo et des régions voisines
(le partie). Ann. Mus. Congo Belge (A) 2: 257-268.

KENSLEY, B. 1974. The status of the Plio-Pleistocene Panopea in southern Africa (Mollusca,
Bivalvia, Hiatellidae). Ann. S. Afr. Mus. 65: 199-215.

KENSLEY, B. 1978. Interaction between coastal processes and lagoonal fauna, between Walvis
Bay and Lideritzbucht, South West Africa. Madoqua 11: 55-60.

KILBURN, R. N. & TANKARD, A. J. 1975. Pleistocene molluscs from the west and south coasts of
the Cape Peninsula, South Africa. Ann. S. Afr. Mus. 67: 183-226.

MiiiarpD, N. A. H. & Scott, K. M. F. 1954. The ecology of South African estuaries. Part VI.
Milnerton Estuary and the Diep River, Cape. Trans. R. Soc. S. Afr. 34: 279-324.

Rocers, J. 1982. Lithostratigraphy of Cenozoic sediments between Cape Town and Elands Bay.
Palaeoecol. Afr. 15: 121-137.

SCHALKE, H. J. W. G. 1973. The Upper Quaternary of the Cape Flats area (Cape Province,
South Africa). Scripta Geol. 15: 1-57.

TANKARD, A. J. 1975a. The marine Neogene Saldanha Formation. Trans. geol. Soc. S. Afr. 78:
257-264.

TANKARD, A. J. 1975b. Thermally anomalous Late Pleistocene molluscs from the south-western
Cape Province, South Africa. Ann. S. Afr. Mus. 69: 17-45.

TANKARD, A. J. 1976. Pleistocene history and coastal morphology of the Ysterfontein—Elands
Bay area, Cape Province. Ann. S. Afr. Mus. 69: 73-119.

VissER, H. N. & Scuocn, A. E. 1973. The geological and mineral resources of the Saldanha Bay
area. Mems geol. Soc. S. Afr. 63: 1-150.

>
.
—
—
~
a

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text

’ 6

en... the Figure depicting C. namacolus...>". ... in Cnamacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

BRIAN KENSLEY

THE FAUNAL DEPOSITS OF A

LATE PLEISTOCENE RAISED BEACH
AT MILNERTON, CAPE PROVINCE,
SOUTH AFRICA

—
ae

ag
VOLUME 95 PART 3 APRIL 1985 ISSN 0303-2515

: SS
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A ‘

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ey NE

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-S1.

THELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
April 1985 April
Part 3 Deel

S -B.9.3:B.9

&

Lp S
S
4 /0UID N NOVI Anes

EARLY PLIOCENE PROCELLARIIFORMES
(AVES)
FROM LANGEBAANWEG,
SOUTH-WESTERN CAPE PROVINCE,
SOUTH AFRICA

By
STORRS L. OLSON

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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EARLY PLIOCENE PROCELLARIIFORMES (AVES) FROM
LANGEBAANWEG, SOUTH-WESTERN CAPE PROVINCE,
SOUTH AFRICA

By
Storrs L. OLSON
Percy FitzPatrick Institute, University of Cape Town*

(With 7 figures and 5 tables)

[MS accepted 14 June 1984]

ABSTRACT

Eight species of sea-birds of the order Procellariiformes are identified among the fossils
collected from early Pliocene deposits at Langebaanweg, south-western Cape Province, South
Africa. All four living families of Procellariiformes are represented, including an albatross
(Diomedeidae), a new species of Oceanites (‘Pelagodroma’, Oceanitidae), five species of
Procellariidae, including three species of Pachyptila, one of which is described as new, and a new
species of diving petrel (Pelecanoididae). At least three of these species appear to have been
breeding in the area, indicating that cold Temperate or Subantarctic oceanic conditions were
present in the south-western Cape in the early Pliocene. The specimens of Oceanites, Pachyptila,
and Pelecanoides provide the first Tertiary records for these genera. Most taxa are very similar to
and perhaps ancestral to living species, with the principal exception of a giant form of Pachyptila
that represents a previously unknown lineage.

CONTENTS

PAGE
MMNGROGUCHOMi a: ste ee Ne hw denn in etn teen rae Jeon 123
SVSUCIMALICS sere neh RICA eres eres Sia a ees tre wecere eee Rave Maw 124
DISCUS SIO Mies eon ee ute lek ae alll Ss a ae ae 141
PNCKnOWIEUSCIMEN(Sta a sere aah Wn ie on mt Aamo stone Ss 144
IRCTOREN CCS Hare eee ewe tara enh nen wot eae re etd Uae tasied 145

INTRODUCTION

Among the abundant fossils of terrestrial and aquatic birds from the early
Pliocene deposits at Langebaanweg (P. Rich 1980) are remains of at least eight
Species representing each of the four families of the strictly marine order
Procellariiformes (albatrosses, storm-petrels, shearwaters, and diving petrels).
The only other site in South Africa from which Tertiary procellariiform fossils
have been recovered is at Duinefontein, also in the south-western Cape and of
approximately equivalent age (Olson in press a). The palaeoceanographic
significance of the marine birds from these early Pliocene faunas is dealt with in a
more general overview (Olson 1983), the scope of the present paper being mainly
systematic.

* Permanent address: National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

128

Ann. S. Afr. Mus. 95 (3), 1985: 123-145, 7 figs, 5 tables.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

The geology and chronology of the Langebaanweg sequence have been
detailed by Hendey (1981a, 19816, 1982). These deposits formed during an early
Pliocene period of marine transgression, under a variety of estuarine and
fluviatile conditions that resulted in the accumulation of remains of both marine
and terrestrial organisms. The Langebaanweg sequence comprises two lithostra-
tigraphic units, the Quartzose Sand Member (QSM) and the Pelletal Phosphorite
Member (PPM). The PPM consists of channel deposits in two different beds, 3aS
and 3aN, the latter of which truncates the former and is thus younger, at least
where palaeontological excavations were undertaken. Both the QSM and PPM
are time-transgressive so that whereas parts of the QSM are older than parts of
the PPM, this relationship does not hold throughout the entire sequence (see
Hendey 19815: 32, fig. 8). There is sufficient evolutionary time represented
between the older and younger portions of the sequence for morphological
changes to have taken place within some species of mammals, e.g. the seal
Homiphoca capensis (De Muizon & Hendey 1980).

Most of the individuals of Procellariiformes come from the QSM deposits
(Table 5), indicating that this unit probably had a somewhat stronger marine
influence than the PPM deposits. The composition of species and individuals of
marine birds and the preservation of the bones themselves contrast with more
typical marine deposits such as at Duinefontein, and indicate that at least the
more abundant species were probably breeding in the vicinity, where they could
have taken advantage of nearby islands that were created by the higher sea levels
of the early Pliocene (Olson 1983).

Fossil specimens described here are in the collections of the division of
Cenozoic Palaeontology of the South African Museum; all fossil specimen
numbers are prefixed by the acronym SAM-PQ, which has been omitted here for
the sake of brevity.

SYSTEMATICS

Order Procellariiformes
Family Diomedeidae
Genus Diomedea Linnaeus, 1758
Diomedea sp.
Material

Proximal end of left tarsometatarsus L12005, from the Quartzose Sand
Member of the Varswater Formation at Langebaanweg.

Discussion

This single bone is the only evidence to date of albatrosses from the Tertiary
of South Africa. The specimen is porous proximally and hence is from a juvenile
individual and may indicate breeding in the vicinity. The proximal width is
17,3 mm, the specimen being from a species approximately the size of the living
Diomedea melanophris.

EARLY PLIOCENE PROCELLARITIFORMES FROM LANGEBAANWEG 125

Although albatrosses today are predominantly birds of southern oceans,
there are but two other Tertiary records from the Southern Hemisphere—a
fragmentary rostrum from the late Miocene of Victoria, Australia, described as
Diomedea thyridata (Wilkinson 1969), and a single toe bone of a larger species
from the early late Miocene near the Valdez Peninsula, Argentina (Olson 1984).
This contrasts with the much better representation of albatrosses in the Northern
Hemisphere where fossils are known from the late Oligocene into the Quaternary
(Olson in press 5).

Family Oceanitidae
Subfamily Oceanitinae

Both on osteological and myological grounds, the storm-petrels fall into two
very distinct groups that are best ranked as subfamilies (see Klemm 1969). The
more specialized of these, to which all the fossils from Langebaanweg clearly
belong, is the Oceanitinae, characterized by very short, stout humeri, ulnae, and
femora, and greatly elongated tibiotarsi and tarsometatarsi. Five genera are
customarily admitted among the species in the Oceanitinae (e.g. Jouanin &
Mougin 1979), but I am unable to discern any osteological basis for considering
either Pelagodroma Reichenbach, 1853, or Garrodia Forbes, 1881, to be distinct
from Oceanites Keyserling & Blasius, 1840. Pelagodroma marina and Garrodia
nereis resemble each other and differ from Oceanites oceanicus and O. gracilis in
having the rostrum longer and more slender and the ridge of bone between
impressions of the supraorbital glands narrower, but neither of these characters
can be regarded as being of generic importance. Hence the first two are included
in Oceanites as O. marinus (Latham) and O. nereis (Gould), respectively.

Within the Oceanitinae, there is an evolutionary trend towards greater size
and increasing specialization of the tarsometatarsus and toes for locomotion
across the surface of the water. In Oceanites this can be described as paddling,
whereas Fregetta and Nesofregetta use the feet to bound rapidly across
the surface, often against strong head winds (D. G. Ainley, Point Reyes Bird
Observatory, pers. comm.). The trend for morphological specialization for such
locomotion reaches its extreme in Nesofregetta fuliginosa, which is the largest
species in the family and in which the distal end of the tarsometatarsus is
expanded and the toes are greatly flattened, being fused by the skin of the web
into a nearly inflexible paddle. The two species of Fregetta are more or less
intermediate in these respects between Oceanites (sensu lato) and Nesofregetta.
Within Oceanites, O. (‘Pelagodroma’) marinus shows perhaps the greatest
tendency towards the specializations of Fregetta and Nesofregetta.

Genus Oceanites Keyserling & Blasius, 1840 (sensu lato)

All the storm-petrel bones from Langebaanweg are referable to a single
species of Oceanitinae that differs from Fregetta in having the tibiotarsus and
tarsometatarsus proportionately longer and much more slender. It differs from

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nesofregetta in lacking the distinctly flattened and expanded distal end of the tarso-
metatarsus and thus agrees with Oceanites in the broad sense as defined above.

Oceanites zaloscarthmus sp. nov.
Figs 1-2

Material

Holotype: L25214, complete right humerus (Fig. 1A), from the Quartzose
Sand Member of the Varswater Formation at Langebaanweg, Cape Province,
South Africa.

Paratypes: In addition to the holotype, 175 other specimens are referred to
this species. These consist of 2 right and 2 left coracoids; 6 complete, 7 proximal,
and 11 distal right humeri; 6 complete, 5 proximal, and 17 distal left humeri;
6 complete and 2 distal right ulnae; 4 complete, 3 proximal, and 4 distal left ulnae;
-2 proximal left carpometacarpi; 4 right and 6 left femora; 14 distal right,
3 proximal left, and 16 distal left tibiotarsi; 2 complete, 17 proximal, and 10 distal
right tarsometatarsi; 1 complete, 13 proximal, and 13 distal left tarsometatarsi. A
list of specimen numbers with exact provenance within the Langebaanweg quarry
is kept at the South African Museum and is also available from the author. The
two carpometacarpi were found in unsorted material and some of the femora
were in amongst the Passeriformes, so additional specimens of the species will
doubtless be found in the material that has already been collected.

Measurements of holotype

Total length 24,80 mm; length from distal end of pectoral crest to distal
extent of dorsal condyle 17,70 mm; width of shaft at midpoint 2,20 mm; distal
width 4,80 mm. (Measurements to nearest 0,05 mm.)

Measurements of paratypes
See Table 1.

Diagnosis

Much larger and more robust than Oceanites oceanicus, O. gracilis, or
O. (‘Garrodia’) nereis. Very similar in size and morphology to O. (‘Pelago-
droma’) marinus but (1) brachial fossa of humerus much shallower and less exten-
sive; (2) olecranon better developed (nearly absent in O. marinus); (3) carpal
tubercle of ulna more pointed, less expanded and less triangular; (4) distal end of
ulna not rotated ventrally; (5) shafts of hindlimb elements more robust; (6) femur
less curved in lateral and medial views; (7) wings of inner and outer trochleae of
tarsometatarsus less prominent and not as expanded to the sides.

Distribution

Early Pliocene Varswater Formation (QSM, PPM 3aN, PPM 3aS) at
Langebaanweg, south-western Cape Province, South Africa.

EARLY PLIOCENE PROCELLARITIFORMES FROM LANGEBAANWEG

TABLE |

Skeletal measurements (to nearest 0,05 mm) of fossil and

living species of Oceanites (‘Pelagodroma’).

CORACOID

Length from head to midpoint
of sternal end

Shaft width at midpoint

HUMERUS

Total length

Length from distal end of pectoral
crest to dorsal condyle

Shaft width at midpoint

Distal width

ULNA

Total length

Length from distal lip of ventral
cotyla to distal end

Shaft width at midpoint

Distal width

CARPOMETACARPUS
Proximal depth

FEMUR
Total length

Proximal width

Shaft width at midpoint
Distal width

TIBIOTARSUS

Distance from proximal articular
surface to distal end of
fibular crest

Distal width

TARSOMETATARSUS

Total length

Proximal width

Shaft width at midpoint
Distal width

O. zaloscarthmus sp. nov.

n

3)
29
14
24

range

13,35-13,60
1,60-1,95

24 ,80—26,50

17,55—19,65
1,90—2,50
4 2)—),29

22,60—23,50

20,95—21,95
1,75—2,20
3,20-3,85

4 ,85—4,90

16,20-—17,50
4,10—4,40
1,70—1,90
3515-9595

11F55—13580
3,59—3,85

40,40-—41,80
3,90-4,55
1,70—2,10
4,25—4,80

* Two skeletons from Peru and one from South Africa.

Etymology

mean

40,90
4.35
1,85
4,45

127)

O. marinus (n = 3*)

range

13,80—14,00
1,45-1,70

DADS ples

17,50-18,55
1,95—2,10
4,60-4,75

22,10—22,95

20,45-21,45
1,80—1,90
3,30=3,55

4,90-—5,00

17,00—17,25
4,00
1,65
3,70—4,20

12,65-14,05
3;50—3;5605

39,05—40,45
4,20-4,35
1,60-1,67
4,45—4,50

mean

4,95

39595
4,30
1,65
4.50

Greek zale, surging sea, and skarthmos, skipping; essentially a rephrasing of

the word pelagodroma.

Remarks

Except for the distinguishing features of the humerus and ulna, the
differences between Oceanites zaloscarthmus and O. marinus are very minor and

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B | C D

Fig. 1. Wing elements of Oceanites (‘Pelagodroma’). A. O. zaloscarthmus sp. nov.,

holotype, L25214, right humerus, cranial aspect. B.O. marinus, USNM 496760, right

humerus, cranial aspect. C. O. zaloscarthmus sp. nov., L21994, left ulna, cranial aspect.
D. O. marinus, USNM 496760, left ulna, cranial aspect. All figures x 3.

it is likely that the Langebaanweg species is ancestral to the living form. A more
specialized condition in the latter is the curious rotation of the distal end of the
ulna ventrally (Fig. 1C, D) with respect to the ‘standard anatomical position’
(Baumel 1979: 5). This is best appreciated when the ulnae are viewed resting on
the caudal (trailing) surface (i.e. with the secondary papillae downward—see
Fig. 1C, D). The deepening of the brachial fossa of the humerus and the
reduction of the olecranon in O. marinus may possibly be correlated with the
ventral rotation of the distal end of the ulna.

The fossils from Langebaanweg constitute only the fourth reported Tertiary
occurrence of the family Oceanitidae. The others consist of two specimens from
the late Miocene of California referred to the genus Oceanodroma (see Howard

EARLY PLIOCENE PROCELLARITIFORMES FROM LANGEBAANWEG 129

Fig. 2. Hindlimb elements of Oceanites (‘Pelagodroma’).
A-B. O. zaloscarthmus sp. nov., L24405, L24390U, left femora in
cranial aspect showing variation in size and robustness. C. O. marinus,
USNM 496760, left femur, cranial aspect. D. O. zaloscarthmus
sp. nov., L24386Q, right tarsometatarsus, plantar aspect. E. O. marinus,
USNM 496760, right tarsometatarsus, plantar aspect. All figures x 2.

1978), and the shaft of a humerus the size and shape of that of Oceanites oceanicus
from Duinefontein (Olson in press a). Despite the relative abundance of
Oceanites zaloscarthmus at Langebaanweg, the species is entirely absent at
Duinefontein. This may reflect differences in the depositional environment at
these two sites. That so many bones of O. zaloscarthmus were concentrated at
Langebaanweg is an almost certain indication that the species was breeding close
by, probably on the islands lying immediately offshore from the Langebaanweg
site (Hendey 1981b, 1982; Olson 1983, in press a). The probable presence of a
breeding colony is also indicated by incompletely ossified bones of young, though
possibly volant, individuals among the fossil sample.

The living species Oceanites marinus is rather widely distributed, nesting on
islands in waters that lie in warm Subantarctic and especially in cool Subtropical
waters. Although the species disperses widely in the non-breeding season, it has
not as yet been recorded from South African waters (Clancey 1980; Harrison
1983). There are breeding populations on islands around western and southern
Australia, in the New Zealand region, and in the Atlantic in the Salvages, Cape
Verdes, and Tristan da Cunha (Jouanin & Mougin 1979). The species was once

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

abundant at St. Helena, probably until after the arrival of man in the sixteenth
century (Olson 1975), and bones are also known from Madeira and Porto Santo
(Harald Pieper, Zoologisches Museum, Kiel, pers. comm.). The species
disappeared in relatively recent times from Amsterdam Island in the Indian
Ocean (Jouanin & Paulian 1960), probably as the result of introduced predators
(Murphy & Irving 1951). The absence of O. marinus in the Benguela Current off
South Africa is the more curious considering that the species is quite abundant in
the Peru Current off South America, which would seem to present similar
conditions. This might be due in part to the extirpation of the population on
St. Helena, although one might expect birds from Tristan da Cunha off South
Africa as well.

Family Procellariidae
Genus Pachypitila Illiger, 1811

The prions (Pachyptila) are unique among the Procellariidae in having the
bill greatly expanded and equipped with lamellae for filtering small prey items.
The tongue and hyoid apparatus are correspondingly enlarged and housed in a
distensible gular sac. With the exception of the very tip of a rostrum, however,
cranial elements of Pachyptila have not yet been identified from Langebaanweg
and fossils from there are assigned to this genus on the basis of characters of the
humerus, which in Pachyptila has a short, blunt ectepicondylar spur (processus
supracondylaris dorsalis) in combination with a very deep brachial fossa, terete
shaft, and lack of expansion of the ventral epicondylar area. The humerus of
Pachyptila is most similar to that of Halobaena, but in that genus the ventral
epicondylar area is slightly expanded and the ectepicondylar spur is deeper
proximo-distally.

Pachyptila salax sp. nov.
Figs 3-5, 6A
Material

Holotype: L25187, complete left humerus (Figs 3A, 6A) from the Quartzose
Sand Member of the Varswater Formation at Langebaanweg, Cape Province,
South Africa.

Paratypes: In addition to the holotype, 202 other specimens are referred to
this species. These consist of 3 complete, 1 scapular, and 1 sternal ends of right
coracoids; 8 complete, 4 scapular, and 1 sternal ends of left coracoids; 1 nearly
complete, 14 proximal, and 16 distal right humeri; 1 nearly complete, 7 proximal,
and 13 distal left humeri; 4 proximal and 4 distal right ulnae; 2 complete,
2 proximal, and 4 distal left ulnae; 3 proximal and 1 distal right carpometacarpi;
2 complete and 2 proximal left carpometacarpi; 1 complete and 3 proximal right
femora; 3 complete, 3 proximal, and 1 distal left femora; 11 right and 5 left distal
ends of tibiotarsi; 7 complete, 13 proximal, and 7 distal right tarsometatarsi;
5 complete, 9 proximal, and 8 distal left tarsometatarsi. With less certainty,

EARLY PLIOCENE PROCELLARIIFORMES FROM LANGEBAANWEG 131

A B C D

Fig. 3. Wing elements of Pachyptila. A. P. salax sp. nov., holotype, L25187, left

humerus, caudal aspect. B. P. vittata, SAM-—ZO56746, left humerus, caudal

aspect. C. P. salax sp. nov., L22224, left ulna, dorsal aspect. D. P. vittata,
SAM-ZO56746, left ulna, dorsal aspect. All figures x 2.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

4 scapulae, 2 fragments of sterna, the distal end of a radius, and 25 pedal
phalanges are assigned to this species on the basis of size. A list of specimen
numbers with exact provenance within the Langebaanweg quarry is kept at the
South African Museum and is also available from the author.

Measurements of holotype

Total length 73,1 mm; length from head to proximal lip of brachial fossa
62,5 mm; length from distal end of pectoral crest to distal end of dorsal condyle
54,7 mm; proximal width through dorsal and ventral tubercles 12,7 mm; width
and depth of shaft at midpoint 4,3 and 3,7 mm; distal width 9,3 mm.

Measurements of paratypes
See Tables 2 and 3.

Diagnosis

Much larger than any known species of Pachyptila (Table 2). Apart from
size, there are few postcranial characters that will distinguish between the species
of Pachyptila, although in P. salax the carpal tubercle of the ulna appears less
pointed and slightly more proximally situated, the alular metacarpal and extensor
process are more perpendicular to the shaft rather than slanting proximally, and
the shaft of the tarsometatarsus appears to be relatively stouter.

Distribution

Early Pliocene Varswater Formation (QSM, PPM 3aN, PPM 3aS, and
Duynefontyn Members) at Langebaanweg and Duinefontein, south-western
Cape Province, South Africa.

Etymology

Greek salax, a sieve, in allusion to the filtering apparatus characteristic of the
living members of this genus. The name is a masculine noun in apposition and
there is no implied allusion to the pejorative Latin adjective of the same
orthography.

Remarks

Pachyptila salax is so much larger than any of the living taxa in the genus
(Table 2), regardless of their status, that there can be no question of its specific
distinctness. It was a giant among prions and adds an entirely new dimension to
our concept of radiation within Pachyptila. Although prions the size of P. salax
may have been endemic to South Africa, it seems likely that birds of this size class
would have been more widely distributed in the past and have become extinct
everywhere since the early Pliocene.

It is unfortunate that there is not more of the bill known for P. salax. The tip
of a rostrum that is tentatively referred to this species (Fig. 5A, C) is not sufficient
even to suggest how much the rest of the bill may have been expanded. Compared

133

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134 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Additional measurements (mm) of Pachyptila salax sp. nov.

n range mean

CORACOID
Length from head to medial angle 9 22,4-24,2 2355
HUMERUS
Proximal width through dorsal and

ventral tubercles 6 12,5-13,3 12,9
Length from distal end of pectoral crest

to dorsal condyle 4 52,2-54,7 335)
Length from head to proximal lip of

brachial fossa 3 62,5-65,3 64,2
ULNA
Proximal width i 5,7-6,5 6,2
Distal width 6 5,5-5,9 S/
CARPOMETACARPUS
Proximal depth 3 8,9-9 2 9,1
Proximal width 5 6,9-7,3 well
Distal width y 6,5-6,5 6,5
TIBIOTARSUS
Distal width iW 5,1-5,5 5,4
TARSOMETATARSUS
Proximal width 30 5,8-6,8 6,4
Shaft width at midpoint 19 2,4-3,1 2,6
Distal width ppp 5,6-6,7 6,2

with P. vittata, the rostral tip of P. salax is markedly broader and considerably less
decurved.

On size alone there are at least three species of Pachyptila in the
Langebaanweg fauna and also at Duinefontein (Olson in press a). Pachyptila
salax is larger than any other species in the genus, Pachyptila species B (see p. 138)
is the size of the two largest living taxa, P. vittata and P. salvini, and Pachyptila
species C (see p. 138) falls within the size ranges of the four smaller living taxa
(Table 2). It is possible that more than one species could be included under
Pachyptila species C. The available specimens of Pachyptila species B and C, all
postcranial, are insufficient to determine anything more than that at least two
species are represented. Although these fossils cannot be distinguished from
living taxa, they cannot be assigned to a particular living taxon nor can one

(facing page)

Fig. 4. Skeletal elements of Pachyptila. A. P. salax sp. nov., L28203U, left carpometacarpus,
ventral aspect. B. P. vittata, SAM—ZOS56746, left carpometacarpus, ventral aspect. C. P. salax
sp. nov., L28174C, right coracoid, ventral aspect. D. P. vittata, SAM—ZOS56746, right coracoid,
ventral aspect. E. P. salax sp. nov., L25531, right femur, cranial aspect. FF. P. vittata,
SAM-Z056746, right femur, cranial aspect. G. P. salax sp. nov., L20691M4, left tarsometatar-
sus, cranial aspect. H. P. salax sp. nov., L24397, right tarsometatarsus, cranial aspect (note
variation in robustness of shaft). I. P. vittata, SAM—ZOS56746, right tarsometatarsus, cranial

aspect. All figures x 2.

EARLY PLIOCENE PROCELLARIIFORMES FROM LANGEBAANWEG 135

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Rostral tips (premaxillae) of Pachyptila. A. P. salax

sp. nov., L28857, ventral aspect. C. Same, lateral aspect.

B. P. vittata, SAM—ZO56746, ventral aspect. D. Same, lateral
aspect. All figures x 2.

assume that the fossils actually were conspecific with any taxon now in existence.

The South African occurrences provide the first fossil record to date for the
genus Pachyptila. The concentration of bones of P. salax at Langebaanweg and
the presence of incompletely ossified bones, probably from pre-fledging indi-
viduals, show that at least P. salax, if not the other species of Pachyptila, was
breeding in the vicinity of Langebaanweg. This indicates that Subantarctic or at
least cool Subtropical marine conditions probably existed nearby. Today, most
breeding colonies of Pachyptila are on islands in Antarctic or Subantarctic waters.
The only colonies north of the Subtropial Convergence are at Tristan da Cunha,
St. Paul and Amsterdam islands, and in the northern part of New Zealand, all of
which lie quite near the Subtropical Convergence.

The systematics of the modern species of Pachyptila are quite complex and a
consensus on the number of species that should be recognized has not been
reached. The divergence in views is exemplified by Harper (1980), who continues
to recognize the six species that have commonly been accepted, and Cox (1980),
who advocates reducing the number to three (though his argument leads to only
two species). Consideration of the osteology of Pachyptila makes it doubtful that
the number of species can be as few as three. Were Cox correct, some of the
species of Pachyptila would exhibit a degree of morphological plasticity with few
parallels among sea-birds.

Fleming (1941) attributed much of the speciation process in Pachyptila to the
effects of Quaternary climatic events. Whereas it is likely that some of the
differentiation between populations that has caused problems in assessing
relationships among living taxa may have arisen as late as the Quaternary, we
have seen that considerable divergence and radiation had already taken place in
Pachyptila by the early Pliocene. It seems probable, therefore, that the principal
species lineages in Pachyptila arose prior to the Quaternary.

EARLY PLIOCENE PROCELLARIIFORMES FROM LANGEBAANWEG 259

se

B

Fig. 6. Left humeri (except C, distal end of right humerus) of Pachyptila in cranial aspect.

A. P. salax sp. nov., holotype L25187. B. P. vittata, SAM-—ZO56746. C. Pachyptila

species B, L25577. —D. Pachyptila species C, L24386D, 1. __E. P. desolata, SAM-ZO56324.
All figures x 2.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pachyptila species B
Fig. 6C

Material

Distal end of right humerus, L25577DF; proximal end of right humerus,
L42830D.

Distribution

Early Pliocene Varswater Formation (QSM, PPM 3aS, and Duynefontyn
Members) at Langebaanweg and Duinefontein, south-western Cape Province,
South Africa.

Remarks

This species is the size of Pachyptila vittata or P. salvini, the two largest living
_ members of the genus, and cannot be distinguished from either on the basis of
available material. See remarks under P. salax.

Pachyptila species C
Fig. 6D

Material

Complete left humerus, L24386D, I; distal ends of left humeri, L25776BO,
L25575B; distal end of right humerus, L25579G2.

Distribution

Early Pliocene Varswater Formation (QSM and Duynefontyn Members) at
Langebaanweg and Duinefontein, south-western Cape Province, South Africa.

Remarks

This species is the size of the smaller living taxa of Pachyptila but the material
is not otherwise diagnostic—see remarks under P. salax. Nine pieces of ulnae,
two coracoids, and the distal end of a tibiotarsus are from procellariids too small
for Pachyptila salax. These most likely belong to one or the other of the smaller
species of Pachyptila.

Genus Puffinus Brisson, 1760
Subgenus Puffinus Brisson, 1760
Puffinus sp.
Material

Distal end of right humerus, L25577CF; proximal end of left humerus,
L56198; right coracoid, L25481.

EARLY PLIOCENE PROCELLARIIFORMES FROM LANGEBAANWEG 139

Distribution

Early Pliocene Varswater Formation (QSM, PPM 3aN, and Duynefontyn
Members) at Langebaanweg and Duinefontein, south-western Cape Province,
South Africa.

Remarks

This is ‘Puffinus species B’ of the Duinefontein fauna. It is similar in
morphology to the living short-tailed shearwater, Puffinus tenuirostris, and was
about the same size, although perhaps slightly smaller. It is illustrated and is
discussed more fully elsewhere (Olson in press a).

Procellariidae gen. et sp. indet.
Material
Left coracoid, L28440J.

Distribution

Early Pliocene, Quartzose Sand Member of the Varswater Formation at
Langebaanweg, south-western Cape Province, South Africa.

Remarks

This specimen is from a procellariid larger than Pachyptila salax and smaller
than the preceding species; thus it represents an additional taxon for Langebaan-
weg. The bone differs qualitatively from any of the species of Puffinus and is
compatible in size with the enigmatic species of fulmarine from Duinefontein
(Olson in press a).

Family Pelecanoididae
Genus Pelecanoides Lacépéde, 1799

The flattened alcid-like humerus and the distinctive lateral reflection of the
head of the coracoid make the three bones discussed below unmistakably
referable to the monogeneric family of diving petrels. All three fossils appear to
belong to a single species that differs only in minor details from living species of
Pelecanoides.

Pelecanoides cymatotrypetes sp. nov.
Biga7

Material

Holotype: Complete left humerus, L14564 (Fig. 7A) from the Quartzose
Sand Member of the Varswater Formation at Langebaanweg, Cape Province,
South Africa.

Paratypes: Proximal end of left humerus lacking internal tuberosity,
L28469T; left coracoid lacking parts of procoracoid and sternocoracoidal
processes, L28855.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B GC D F

Fig. 7. Proximal portions of left humeri in caudal aspect (A—C) and left coracoids in ventral

aspect (D, E) of Pelecanoides. A. P. cymatotrypetes sp. nov., holotype, L14564.

B. P. cymatotrypetes sp. nov., L28469T. C. P. urinatrix exsul, USNM 553240.
D. P. cymatotrypetes sp. nov., L28855. E. P. urinatrix exsul, USNM 553242.

Measurements of holotype

Total length 43,2 mm; proximal width 9,2 mm; shaft width and depth at
midpoint 3,4 and 2,1 mm; distal width 7,1 mm.

Measurements of paratypes

See Table 4 for measurements of coracoid.

Diagnosis

Differs from living species in lacking the distally projecting protuberance on
the caudal surface of the head of the humerus; ventral tubercle in ventral view
shorter and deeper. Coracoid with head projecting more ventrally and less
medially than in living species.

Distribution

Early Pliocene, Quartzose Sand Member of the Varswater Formation at
Langebaanweg, south-western Cape Province, South Africa.

Etymology

Greek kyma, wave, and trypetes, borer, from the habit of the living species of
flying straight through the crests of waves. The name is a masculine noun in
apposition.

EARLY PLIOCENE PROCELLARITFORMES FROM LANGEBAANWEG 141

TABLE 4
Measurements (mm) of living and fossil taxa of Pelecanoides.

Coracoid length

Humerus from head to midpoint
Species n length of sternal facet
P. cymatotrypetes sp. nov. 1 43,2 24,6
P. urinatrix exsul 2 43,0; 44,1 PIB) Pop Poet
P. urinatrix chathamensis 1 40,6 21,8
P. urinatrix subsp. (Argentina) 1 42,1 Ws)
P. magellani 2 43,3; 44,8 13) 83 Hs)
P. georgicus 1 3953 21,4
Remarks

This species is very similar to living forms except for the less extensive
ossification of the head of the humerus, in which respect it is probably primitive.
In size it is similar to Pelecanoides urinatrix exsul or P. magellani, but the bones
are stouter than in the latter and the resemblances of the fossil are greatest to
enUENCXS UL.

The three bones of P. cymatotrypetes provide the only Tertiary record of the
Pelecanoididae. A supposed Tertiary occurrence of Pelecanoides in New Zealand
was subsequently shown to be Quaternary in age (T. Rich ef al. 1979). Modern
diving petrels are confined to cold Temperate or Subantarctic waters. None has
been recorded from South Africa (Clancey 1980) and the nearest breeding colony
is at Tristan da Cunha. Although considered to be ‘sedentary’ (Jouanin & Mougin
1979), high densities of diving petrels may occur some 1 300 to 1 600 km from the
nearest land (D. G. Ainley, pers. comm.). They are nevertheless not as vagile as
other members of the order.

DISCUSSION

Despite the fact that at present the order Procellariiformes is far more
diverse in the Southern Hemisphere, its fossil record has hitherto been largely
confined to the Northern Hemisphere, where the taxa represented consist mainly
of albatrosses and a diversity of shearwaters of the genus Puffinus (Olson in
press b). The early Pliocene deposits at Langebaanweg and at Duinefontein
(Olson in press a) thus provide our first important insights into the procellariiform
fauna of the southern oceans in the late Tertiary, as well as the first Tertiary
records for the genera Oceanites, Pachyptila, and Pelecanoides.

The fossils available from deposits in the Northern Hemisphere seem to
indicate that evolution within the Procellariiformes proceeded rather slowly from
the Miocene onward, with species’ lineages persisting for long periods with
relatively little morphological change. Being extremely vagile, procellariiforms

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

adapted to particular oceanic conditions probably moved with them whenever
global climatic changes caused shifts in surface water temperatures or salinities.
Thus these birds have been able to remain with a ‘stable’ environment over long
periods of geological time.

The Benguela Current and the cold upwelling associated with it did not
originate until early late Miocene (Siesser 1980). It is therefore unlikely that a
procellariiform fauna with such characteristically Subantarctic elements as seen at
Langebaanweg and Duinefontein could have become established in South Africa
before then. These species almost certainly did not just appear de novo, however,
and were therefore probably present at higher latitudes during the Miocene and
merely moved into southern Africa when conditions there became suitable
(Olson 1983).

The history of certain marine organisms in South Africa may thus have close
parallels with those on the western coast of South America. Zinsmeister (1978)
has suggested a correlation between the formation of the West Antarctic ice sheet
and the disruption of major current patterns in the Southern Hemisphere, which
sent cold currents up the Pacific coast of South America and permitted cold-water
faunas to expand northward out of the higher latitudes. The situation with South
African sea-birds would appear to agree with Zinsmeister & Feldmann’s (1984)
view of the higher southern latitudes as a centre of origin for numerous animals
that evolved early in the Tertiary but that did not disperse to middle latitudes
until the Neogene.

Of the eight species of Procellariiformes at Langebaanweg (Table 5), six do
not differ in any major way from living species and each could possibly be
ancestral to some existing form. This is certainly true of Oceanites zaloscarthmus
and Pelecanoides cymatotrypetes, which are very likely to be on a direct line with
O. marinus and P. urinatrix, respectively. The material of Diomedea, Puffinus,

TABLE 5

Distribution of bones of Procellariiformes in the various units of the Varswater Formation at

Langebaanweg. Number of specimens in first column, minimum number of individuals in

parentheses. Pedal phalanges tentatively assigned to Pachyptila salax are not included. The
material from PPM 3aS may include some specimens redeposited from the QSM.

QSM PPM 3aS PPM 3aN
Diomedea sp. le ) —_-_ — =
Oceanites zaloscarthmus 147 (35) AD (7) 9 5G)
Pachyptila salax 187 (G3) Si) SG)
Pachyptila species B i Cl) cl) —_—- —
Pachyptila species C 4 (3) —_- — —_- —
Pachyptila spp. B or C Sa) Lele) 2a hy
Procellariidae, gen. & sp. indet. i (a) —- — —_- —
Puffinus sp. 2 (i) —- — | as)
Pelecanoides cymatotrypetes 3 2) —_-_ — =
TOTAL 355 (80) 30 (5) 20 (14)

EARLY PLIOCENE PROCELLARIIFORMES FROM LANGEBAANWEG 143

and the two smaller species of Pachyptila is too incomplete for confident
assessment, but nevertheless presents nothing to suggest that any of these taxa
represent lineages with no living descendants.

The coracoid of the unidentified genus of Procellariidae, if from the same
species as the enigmatic fulmarine from Duinefontein, might indicate an extinct
lineage. The most interesting of the procellariiform taxa at Langebaanweg is
Pachyptila salax, which represents a totally extinct line of giant prions. Why this
largest species in the genus should have become extinct while smaller species have
remained diverse and abundant in the Subantarctic realm is not readily apparent.

Only two of the procellariiforms at Langebaanweg, Oceanites zaloscarthmus
and Pachyptila salax, are abundantly represented and appear to be more than
incidental. Bones of juvenile individuals of both of these species are present in the
Langebaanweg deposits and both were thus probably breeding in the vicinity, as
may also be presumed for Pelecanoides cymatotrypetes because of the more
sedentary nature of the modern members of this genus. The procellariiform fauna
at Langebaanweg, having formed at or near the site of breeding colonies, thus
contrasts with those, such as at Duinefontein and at most localities in the
Northern Hemisphere, where diversity is higher due to the presence of migrant
and wintering species, but numbers of individuals per species are lower.

The great preponderance of specimens and individuals of Procellariiformes
at Langebaanweg is in the QSM rather than in the channel deposits of the PPM
(Table 5). As these are strictly marine taxa, this reflects either a more marine
depositional environment in the QSM or at least an enhanced probability in the
QSM of post-mortem transportation of sea-birds from a more purely marine
situation. The minimum numbers of individuals in Table 5 were calculated by
regarding specimens from each collecting site within the quarry as a separate
sample, which perhaps has the potential of yielding too high a value. Considering
that we are dealing with minima anyway, it is doubtful that this has exaggerated
the number of individuals that contributed bones to the total fossil sample. Under
conditions of terrestrial deposition it was found that the relative abundance of
species of Procellariiformes on St. Helena Island did not differ significantly
whether calculated by total number of specimens or minimum number of
individuals (Olson 1975).

The fossil Procellariiformes and other sea-birds indicate the presence of cold
waters off the Atlantic coast of South Africa in the early Pliocene. Since that
time, Procellariiformes have ceased to breed in South Africa, and, in fact, no
species of this order breeds on any continental African island today. Because the
Benguela Current still provides cold upwelling off the southern African coast, the
cause of the apparent retreat of certain species to higher latitudes and the
extinction of other species, such as Pachyptila salax and several penguins (Olson
1983), is not readily perceived. Certainly the change in size, number, and
character of suitable breeding islands that came about as a result of falling sea-
levels would very likely have had a marked effect on Procellariiformes. This
would be particularly true for burrowing species because the sea-bird islets that

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

remain in the Cape region today are largely devoid of soil in which to burrow.
Nevertheless, many species of Procellariiformes nest exposed on the ground or in
crevices in rocks and would seemingly have been less severely affected by
geomorphological changes in islands.

Not only has there been a nearly complete turnover and restructuring of the
marine avifauna of the Cape region since the early Pliocene (Olson 1983), but the
same holds true for pinnipeds as well. The seal Homiphoca capensis, which has its
nearest affinity with the Antarctic crab-eater seal Lobodon carcinophagus (see
De Muizon & Hendey 1980), was abundant at Langebaanweg in the Pliocene but
since then has become extinct and has been replaced by the fur seal Arctocephalus
pusillus, whose congeners are found in Subantarctic and Subtropical waters as
well as the Antarctic. Such a pattern does not seem explicable solely by changes in
the nature of offshore islands.

Very likely there was a combination of factors responsible for the
pronounced changes in the fauna of marine homeotherms observed since the
early Pliocene in South Africa. Present evidence suggests that oceanographic
conditions have not remained stable and have become less advantageous for
organisms that are now characteristic of colder waters at higher latitudes.

ACKNOWLEDGEMENTS

My research on South African fossils was instigated and supported by the
Percy FitzPatrick Institute of African Ornithology, University of Cape Town, in
which connection I especially thank Timothy M. Crowe. I am also grateful for
additional funds received from the University of Cape Town, the Council for
Scientific and Industrial Research, Pretoria, and the Smithsonian Institution. The
South African Museum graciously provided office space and much logistic
support; Philippa Haarhoff deserves special mention for attending to many of my
research needs, including providing data for minimum numbers of individuals and
double-checking specimen numbers. My studies have benefited greatly from
information and advice supplied by Q. B. Hendey and from discussions with
G. Avery, R. K. Brooke, J. Cooper, D. C. Duffy, P. Haarhoff, P. A. R. Hockey,
H. F. James, and R. P. Prys-Jones. Modern comparative material examined came
mainly from the zoological collections of the South African Museum (prefixed by
SAM-—ZO) and the National Museum of Natural History, Smithsonian Institution
(USNM), supplemented by specimens from the Transvaal Museum kindly lent by
A. C. Kemp. Clive Booth (South African Museum) deserves much credit for his
labours in providing the photographs. I am grateful to Helen F. James for
assistance in several aspects of this study and much useful discussion of the
manuscript, which was also read by David G. Ainley, Richard K. Brooke,
Kenneth E. Campbell, David C. Duffy, Philippa Haarhoff, Hildegarde Howard,
Christian Jouanin, and David W. Steadman.

EARLY PLIOCENE PROCELLARITIFORMES FROM LANGEBAANWEG 145

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CLANCEY, P. A. ed. 1980. $.A.O.S. checklist of southern African birds. Johannesburg: Southern
African Ornithological Society.

Cox, J. B. 1980. Some remarks on the breeding distribution and taxonomy of the prions
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FLEMING, C. A. 1941. The phylogeny of prions. Emu 41: 134-155.

Harper, P. C. 1980. The field identification and distribution of the prions (genus Pachyptila),
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—

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

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Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text _
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(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
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(c) Scientific names, but not their vernacular derivatives
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Punctuation should be loose, omitting all not strictly necessary

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Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

STORRS L. OLSON

EARLY PLIOCENE
PROCELLARIIFORMES (AVES)
FROM LANGEBAANWEG,
SOUTH-WESTERN CAPE PROVINCE,
SOUTH AFRICA

=
VOLUME 95 PART 4 _—APRIL_ 1985 ISSN 0303-2515

CAPE TOWN

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BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIsCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THEELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

PN NALSYOFR THE SOUTHOAFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
April 1985 April
Part 4 Deel

AN EARLY PLIOCENE MARINE AVIFAUNA
FROM DUINEFONTEIN,
CAPE PROVINCE, SOUTH AFRICA

By
STORRS L. OLSON

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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AN EARLY PLIOCENE MARINE AVIFAUNA FROM
DUINEFONTEIN, CAPE PROVINCE, SOUTH AFRICA

By
Storrs L. OLSON

Percy FitzPatrick Institute, University of Cape Town*

(With 3 figures and 1 table)

[MS accepted 14 June 1984]

ABSTRACT

Late Tertiary marine deposits of the Varswater Formation at Duinefontein, Cape Province,
South Africa, have yielded remains of 16 or 17 species of sea-birds (Sphenisciformes,
Procellariiformes, Pelecaniformes) and one land-bird (Galliformes, Phasianidae). Most of the
sea-birds are characteristic of cold waters, indicating that these deposits are probably no older
than late Miocene, the age of origin of the Benguela upwelling, and the species composition of
the marine avifauna correlates well with nearby early Pliocene deposits at Langebaanweg.
Differences between the sea-bird faunas at these two sites may be related to differences in the
depositional environments.

CONTENTS

PAGE
| (DUOC LEK MOTT Nese oe ie a0 ae eee oes ear eS 147

Synopsis of the geology and age of Tertiary sea-bird sites in
Be @ape PLOVINGCEs Berets 2 abet as cathe nants die ae eee eee Sires = 148
SVSc MRA TI CSUSee mes rasicg ace hentai eS ond nn owe ea ne iby
DISCUSSIONS reat oe ees yee nee Seo ers CO eee 161
FNCKHOWICUSCIICIILS set None: utter es Ne tee eeia ee 163
IRCKC TEMG Sirs ae a eas ay vats she naa Heisler ee nce aaa 163

INTRODUCTION

During the construction of the Koeberg nuclear power station in 1978,
fossiliferous sediments were exposed in two excavations at Duinefontein farm on
the coast 30 km north of Cape Town. Vertebrates recovered here consist of
marine birds, cetaceans, sharks, and bony fishes, as well as a small terrestrial
component including ungulate, lagomorph, snake, and turtle remains (Rogers
1979). To date, the only study of any of the vertebrates from this site is Simpson’s
(1979b) report on six penguin bones, identified as belonging to two species, only
one of which, Nucleornis insolitus, was represented by sufficiently diagnostic
material to merit naming. Since then, many more avian fossils have been
obtained, so that now there are over 70 reasonably diagnostic bones of penguins
from at least four species, as well as specimens assignable to 12 or 13 species of

* Permanent address: National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

147

Ann. S. Afr. Mus. 95 (4), 1985: 147-164, 3 figs, 1 table.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Procellariiformes and Pelecaniformes, and a single fragment of a francolin
bone (Phasianidae, Galliformes), the last being the only terrestrial bird in the
fauna.

Although many of the specimens are very fragmentary and can be identified
only tentatively, the fauna nevertheless provides important new information on
the distribution and occurrence of sea-birds in the late Tertiary of the South
Atlantic. All fossil specimens are in the collections of the section of Cenozoic
Palaeontology at the South African Museum and specimen numbers are preceded
by the acronym SAM-PQ, here omitted for brevity.

SYNOPSIS OF THE GEOLOGY AND AGE OF
TERTIARY SEA-BIRD SITES IN THE CAPE PROVINCE

Tertiary sea-birds have been obtained from three localities (Fig. 1) in the
south-western Cape Province—Duinefontein, Ysterplaat, and Langebaanweg
(Olson 1983). As the first two of these sites have in the past been attributed
incorrectly to the Miocene (Simpson 1973, 19795), it is appropriate to review here
some of the new stratigraphic and faunal information altering that interpretation.
Simpson, of course, is blameless in referring to the Duinefontein and Ysterplaat
sites as Miocene, for he relied entirely on preliminary assessments that had been
communicated to him. Nevertheless, the belief that the penguin fossils from these
sites were Miocene in age probably affected his taxonomic conclusions to some
degree.

The best known of the above sites is Langebaanweg, the stratigraphy and
mammalian fauna of which has been exhaustively treated by Hendey and others
in numerous papers (see Hendey 1981a, 1981b, 1982, and references therein).
Virtually all sea-bird fossils from Langebaanweg come from two extremely
fossiliferous units of the Varswater Formation that are early Pliocene (5 Ma) in
age (Hendey 1981a, 1981b, 1982). Fossils were deposited under a variety of
estuarine, palustrine, and fluviatile conditions. The vertebrates, although
dominated by terrestrial forms, have a strong marine component that includes
sharks, seals, and whales, as well as sea-birds. The marine avifauna consists for
the most part of numerous individuals of relatively few species that probably bred
on nearby islands (Olson 1983, 1985). .

In the revised view of the stratigraphy of the Duinefontein sediments, the
beds containing fossil birds are now considered to belong to the Duynefontyn
Member of the Varswater Formation (Dingle e/ al. 1983, modified from Rogers
1979). The bird remains were apparently deposited during the same early
Pliocene marine transgression during which the Langebaanweg deposits were
formed. The deposits are 8,2 to 8,5 m below present sea level. The environmental
setting at the time of deposition is thought to have been a lagoon sheltered by a
barrier spit that was breached by storm or spring tides (Rogers 1979). This
interpretation accords with the fact that some of the bird bones are fairly well
preserved, whereas others are heavily worn. The avifauna consists almost entirely

EARLY PLIOCENE MARINE AVIFAUNA 149

IO ae

St Helena
Bay

34°

18°30

Fig. 1. Tertiary sea-bird localities in the south-western Cape Province showing
their relationship to possible shoreline configuration (shaded portion) in the early
Pliocene (modified from Olson 1983).

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

of pelagic species, a number of which may have been non-breeding migrants. This
indicates direct access to the open ocean, at least at times. Of the 16 or 17 species
in the Duinefontein fauna, 10 to 12 appear to be shared with Langebaanweg,
which is further evidence of their probable contemporaneity.

The deposits at Ysterplaat Air Force Base, on the outskirts of Cape Town,
are at 10 m above sea level and 1,5 km from the present coastline. Tankard
(1975a, 1975b) assigned the Ysterplaat deposits to his Saldanha Formation, the
type section of which is much farther north on Hoedjiespunt in Saldanha Bay.
This formation was erected by Tankard ‘as a convenience to accommodate all
phosphate rock-bearing horizons in the western Cape’, the assumption at the time
being that all such phosphate rocks were Miocene in age, whereas subsequently ‘it
has been shown that thick phosphate rock units occur in the upper part of the
Varswater Formation’ (Dingle et al. 1979: 91). Consequently, ‘the original
definition of the Saldanha Formation, as a lithostratigraphic unit distinct from the
Varswater Formation, cannot be demonstrated with present data, and .. . the
use of the term ‘Saldanha Formation” [should] be discontinued’ (Dingle et al.
1979: 81). Because the assumption that phosphatic rocks must be Miocene was
erroneous, because there was no real basis for assigning the Ysterplaat deposits to
the Saldanha Formation in the first place, and because the existence of such a
formation cannot be demonstrated, there is no basis for considering the
Ysterplaat deposits to be Miocene in age. These deposits appear to be purely
marine in origin, the only vertebrate fossils present being those of penguins,
whales, and sharks. The penguin material is in very poor condition, although part
of it provided the basis for the species Simpson (1973) named ?Palaeospheniscus
huxleyorum. As far as the material permits, the three species of penguins at
Ysterplaat appear to be the same as the three largest species common to both
Duinefontein and Langebaanweg. There is no evidence at present that the
Ysterplaat fossils are not of approximately equivalent age.

Thus, new stratigraphic revisions, as well as the nature of the marine
avifauna, indicate that all three sites in the south-western Cape from which fossil
sea-birds have been recovered are likely to be early Pliocene in age, the deposits
all probably having formed at some phase of the same sea-level cycle. On the
basis of molluscs from the Gravel Member underlying the Varswater Formation
at Langebaanweg, Hendey (1981a, 1981b) postulated that late Miocene marine
temperatures were warmer than during the deposition of the succeeding early
Pliocene sediments. Furthermore, Siesser (1980) has shown that the Benguela
Current and its associated cold upwelling did not originate off the south-western
coast of Africa until the early late Miocene. The marine avifaunas from
Duinefontein and Langebaanweg contain a number of Subantarctic, cold-water
species of Procellariiformes as well as a diversity of penguins (Olson 1982, 1985,
in prep.). Such assemblages would have been unlikely to have been present prior
to the origin of the colder waters and increased marine productivity that the
Benguela upwelling would have provided; hence these fossils would not be older
than late Miocene in any case.

EARLY PLIOCENE MARINE AVIFAUNA 154

SYSTEMATICS

Order SPHENISCIFORMES
Family Spheniscidae

There are at least four species of penguins from Duinefontein, corresponding
to the four species named from South Africa by Simpson (1971, 1973, 1975,
1979b). These species, however, were described in four different extinct genera,
whereas it is now believed that all probably belong to a single genus (Olson in
prep.). This genus is either distinct from all living genera but closely related to
Spheniscus, or the South African fossil penguins are actually primitive forms of
Spheniscus and should be referred to that genus. In the former case, the generic
name /nguza Simpson (1979a) would apply. A decision on the generic status of
these penguins would perhaps be facilitated by examination of early Pliocene
penguins from Peru (see De Muizon 1980, 1981). Rather than creating new
combinations at this point, each species has been listed in the genus in which it
was last placed by Simpson, with the generic name in quotes to indicate present
uncertainties.

‘Nucleornis’ insolitus Simpson, 1979

Material

Holotype: right tarsometatarsus, MBD4. Paratype: right tarsometatarsus,
MBD3. Specimens referred herein: worn proximal end of left coracoid, MBD215;
distal end of left radius, MBD399; proximal ends of right radii, MBD7, MBD 161;
distal end of right ulna, MBD304 + MBD160; shafts of left femora, MBD318,
MBD532; shaft of right tibiotarsus, MBD320; worn metatarsal (probably R4),
MBD219. Minimum number of individuals, 2.

Remarks )

This species is the largest of the South African fossil penguins and is larger
than any living penguin except the two species of Aptenodytes. It is the only
species for which Duinefontein is the type-locality. The two tarsometatarsi
studied by Simpson (1979a) remain the only really diagnostic specimens, although
a few others from Duinefontein, Ysterplaat, and Langebaanweg are assigned to
this species on size alone.

‘Dege’ hendeyi Simpson, 1975
Material

Worn right radius lacking distal end, MBD303; shaft of left tibiotarsus,
MBD533.

Remarks

This rare species is intermediate in size between “Nucleornis’ insolitus and
‘? Palaeospheniscus’ huxleyorum. The only reasonably diagnostic material is from

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Langebaanweg, the type-locality, with fragmentary specimens from Duinefontein
and Ysterplaat being referred on size.

‘? Palaeospheniscus’ huxleyorum Simpson, 1973

Material

Fragment of right mandibular ramus, MBD201; right clavicle, MBD419
[possibly too large for this species]; fragmentary right coracoids including at least
the glenoid area, MBD153, MBD154, MBD214, MBD310, MBD313, MBD528;
fragmentary left coracoids including at least the glenoid area, MBD2, MBD152,
MBD308, MBD311, MBD312; sternal ends of left coracoids, MBD203,
MBD307; proximal ends of right humeri, MBD296, MBD527; shafts of right
humeri, MBD207, MBD211, MBD297; distal ends of right humeri, MBD210,
MBD294; complete left humerus, MBD151; complete left ulna, MBD202;
complete right radius, MBD302; proximal end of right radius, MBD129A;; left
radius lacking distal end, MBD305; distal end of left radius, MBD129B; ulnare,
MBD418; alar phalanx, MBD314; complete right femur, MBD92; proximal ends
of right femora, MBD300, MBD301; distal ends of right femora, MBD157,
MBD315, MBD400, MBD471; shafts of right femora, MBD218, MBDS535;
fragmentary tibiotarsi, MBD212, MBD213, MBD316, MBD515; complete left
tarsometatarsus, MBD292; shaft of left tarsometatarsus, MBD468. Minimum
number of individuals, 6.

Remarks

This species is somewhat larger than the largest individuals of the living
South African penguin Spheniscus demersus. Originally described from Yster-
plaat, it is the most abundant penguin at Duinefontein and the second most
abundant at Langebaanweg, where most of the material was incorrectly
attributed to the larger species ‘Dege’ hendeyi by Simpson (1975).

‘Inguza’ predemersus (Simpson, 1971)

Material

Right mandibular articulation, MBD322; right quadrate, MBD342; scapular
end of left coracoid, MBD155; complete right humerus, MBD295; shaft of right
humerus, MBD10; shafts of left humeri, MBD11, MBD70; distal end of left
humerus, MBD530; complete left ulna, MBD159; right radius, MBD204;
complete right carpometacarpus, MBD293; shaft of right tibiotarsus, MBD319;
left tibiotarsus lacking proximal end, MBD1; shaft of left tarsometatarsus lacking
fourth metatarsal, MBD487. Minimum number of individuals, 3.

Remarks

This is the commonest penguin at Langebaanweg, the type-locality, and is
the second most abundant penguin at Duinefontein, although it was not

EARLY PLIOCENE MARINE AVIFAUNA 153

recovered at Ysterplaat. It is a small species, somewhat smaller than Spheniscus
demersus, to which it was originally thought to be ancestral (Simpson 1971).

In addition to numerous unidentifiable scraps of penguin bone, there are
specimens that seem to be too large for ‘Inguza’ predemersus and too small for
‘?Palaeospheniscus’ huxleyorum: fragmentary right coracoids, MBD309,
MBD401; shaft of right femur, MBD317; distal end of right femur, MBD534:
shaft of right tibiotarsus, MBD158: distal end of left tibiotarsus, MBD298. These
fossils cannot be assigned positively, nor can it be stated with certainty that they
represent a fifth species. Some bones from Langebaanweg are also of this size.

Order PROCELLARIIFORMES
Family Oceanitidae
Subfamily Oceanitinae
Oceanites sp.

Material
Shaft of right humerus with distal portion of scar for M. pectoralis, MBD260.

Remarks

Although very fragmentary, this specimen comes from a species much
smaller than any other bird known from Duinefontein and is sufficiently
diagnostic for assignment to the short-winged subfamily Oceanitinae of the
Oceanitidae. It is from a bird smaller than Oceanites zaloscarthmus, a species
common at Langebaanweg (Olson 1985), and agrees in size and details with the
living species Oceanites oceanicus.

Family Procellariidae
Fulmarinae, gen. et sp. indet.
Fi

4

gg

Material
Distal end of right humerus, MBD334.

Remarks

This is one of the better-preserved specimens from the Duinefontein site and
is quite singular in its morphology. The very deep brachial fossa and triangular,
proximally pointing ectepicondylar spur (processus supracondylaris dorsalis) give
it a strong superficial resemblance to a gull (Laridae); yet the lack of distinct
tricipital sulci, the more expanded and rounded ventral epicondylar area, and the
heavier shaft in ventral view show that it cannot be a gull and must belong in the
Procellariidae. The shortness of the ectepicondylar spur, the deep brachial fossa,
and the less flattened and expanded ventral epicondylar area suggest that this bird
belongs with the ‘fulmarine’ group of petrels, rather than with Procellaria,
Calonectris, or Puffinus. In size it is intermediate between the smaller Daption on

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

B

Fig. 2. Distal end of right humerus of Fulmarinae,
gen. et sp. indet., MBD334. A. Caudal aspect.
B. Cranial aspect. X 2.

the one hand, and the larger Fulmarus—Thalassoica on the other. Of these
genera, it is more similar to Daption in not having the brachial depression
extending as far proximally. It differs from these genera, as well as from
Pagodroma, Halobaena, and Pachyptila, in the shape of the ectepicondylar spur
and in the deeper brachial fossa. If correctly referred to the fulmarine petrels, this
specimen would represent a species in a size-class that has become extinct. A
coracoid from Langebaanweg may also be referable to this species because it is
too large for the largest species of Pachyptila yet is not referable to the genus
Puffinus or any of its close relatives (Olson 1985).

Pachyptila salax Olson, 1985

Material

Incomplete distal end of right humerus, MBD261; incomplete distal end of
left humerus, MBD253; pieces of shaft of right humerus, MBD149, MBD246;
proximal end of left carpometacarpus, MBD463; distal end of left carpometacar-
pus, MBD237; fragment of right coracoid, MBD387; proximal end of left
tarsometatarsus, MBD410. Minimum number of individuals, 2.

Remarks

The specimens listed here belong to a species smaller than any of the other
procellariids in the fauna except the two following. The only reasonably

EARLY PLIOCENE MARINE AVIFAUNA 155

diagnostic specimen is MBD261, which, although lacking the condyles, has the
ectepicondylar spur and part of the brachial fossa remaining. The spur is short
and rounded and the brachial fossa is deep, as in Pachyptila, and the specimen
agrees in size and other details with the giant species Pachyptila salax that
dominates the procellariiform fauna at Langebaanweg (Olson 1985). The other
material from Duinefontein is referred to this species solely on the basis of size.

Pachyptila species B

Material
Distal end of left humerus, MBD546.

Remarks

This specimen is from a species of Pachyptila the size of P. vittata, the largest
of the living species of the genus. It is indistinguishable from a comparable
specimen from Langebaanweg (Olson 1985).

Pachyptila species C

Material

Right coracoid, MBD322; shaft of left humerus, MBD464; shaft of right
humerus, MBD339.

Remarks

This species is smaller than P. vittata and is similar in size to the smaller living
species P. desolata, being smaller than any of the other procellariids in the
Duinefontein fauna. The coracoid is the only reasonably well-preserved
specimen, the others being included only on size. This species would be of the
same size as the smallest species of Pachyptila from Langebaanweg.

Procellaria sp.

Material

Shaft of left humerus with most distal part of pectoral crest and scar for
M. pectoralis, MBD86 + 86C.

Remarks

This specimen is assigned to the genus Procellaria (including Adamastor) on
size alone, the members of this genus being much smaller than Macronectes and
markedly larger than any of the other forms of Procellartidae. The fossil also
agrees with Procellaria in the pronounced, wide distal scar for M. pectoralis. This
is the first Tertiary record of the genus Procellaria.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Calonectris sp.

Material

Right carpometacarpus lacking minor metacarpal and distal end, MBD144;
worn distal end of right tarsometatarsus, MBD324.

Remarks

These specimens are from a large shearwater slightly larger than Calonectris
diomedea or Puffinus gravis. The alular metacarpal is decidedly notched and thus
very unlike Fulmarus, Daption, or Macronectes. In size and robustness of shaft,
MBD144 is most similar to Calonectris diomedea, although the pisiform process is
more reduced in the fossil.

Puffinus (Puffinus) species A
Fig. 3A

Material

Complete right ulna, MBDS545; shaft of right homerus, MBD564; shaft of left
humerus, MBD336.

Remarks

The ulna cited here is the most complete of the diagnostic procellariiform
fossils from Duinefontein, lacking only part of the olecranon. The shaft is very
short, thick, and curved compared to the ulnae in modern species of Puffinus of
comparable size. The distal portion of a shaft of a humerus is tentatively assigned
here as it is much compressed but larger than in either of the other two species of
Puffinus from Duinefontein.

Puffinus (Puffinus) species B
Fig. 3B, C

Material

Distal end of left humerus, MBD12; shafts of left humeri, MBD407,
MBD496; distal ends of right humeri, MBD146, MBD244, MBD497; shafts of
right humeri, MBD565, MBD407; distal end of left tibiotarsus, MBD458; distal
end of left tarsometatarsus, MBD13. Minimum number of individuals, 2.

Remarks

This species has the very flattened humerus characteristic of the subgenus
Puffinus. It is nearest to the living species P. tenuirostris in size and morphology
but appears to be slightly smaller. A few bones from Langebaanweg (e.g. Fig. 3B)
have also been assigned to this species (Olson 1985).

EARLY PLIOCENE MARINE AVIFAUNA 157

Fig. 3. Wing elements of Puffinus. A. Puffinus species A, MBDS545, right ulna, ventral

aspect. B-D. Distal ends of right humeri, cranial aspect. B. Puffinus species B, L25577F

(Langebaanweg). C. Puffinus species B, MBD244 (Duinefontein). D. Puffinus species C,
MBD337. All figures x 2.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Puffinus (Puffinus) species C
Fig. 3D

Material

Distal end of right humerus, MBD337; shaft of right humerus, MBD86; shaft
of right ulna, MBD566; proximal end of left ulna, MBD541. Minimum number of
individuals, 2.

Remarks

These fossils appear to be from a species of Puffinus smaller than Puffinus
(P.) species B and similar in size to P. p. puffinus but not having the shaft as
compressed or the brachial depression as reduced as in that species. Morphologi-
cally it is thus more like a small version of P. tenuirostris.

Medium-sized indeterminate Procellariidae

Among the remaining specimens from Duinefontein are 13 fragments of
humeri, 5 of ulnae, 6 of carpometacarpi, 4 scapulae, 1 coracoid, and 6 basal
phalanges of the major digit of the wing that can be assigned to the Procellariidae.
Among the species recognized from Duinefontein, these specimens are too large
for any Pachyptila and too small for Calonectris or Procellaria, but they are not
otherwise sufficiently diagnostic to be assigned to any of the four medium-sized
species recognized here, or to permit the recognition of any additional species.

Order PELECANIFORMES

Family Sulidae
Sula sp.

Material
Proximal half of phalanx 1 of major digit of wing, MBD340.

Remarks

This specimen comes from a sulid much smaller than the living Cape gannet
Morus capensis and is thus likely to be referable to the same small species of Sula
that is known from Langebaanweg (Olson 1983). The material from Langebaan-
weg is so scanty that it does not merit detailed treatment in a separate publication,
and therefore will be dealt with here.

The four specimens of Sula sp. from Langebaanweg consist of two sternal
ends of right coracoids from the Pelletal Phosphorite Member (Bed 3aS) and the
scapular end of a coracoid and distal end of a humerus from the Quartzose Sand
Member. These represent a minimum of four individuals. Measurements of these
specimens are as follows: coracoid—head to sternal lip of glenoid facet 20,1 mm,
length and width of glenoid facet 10,8 x 6,5 mm, length and width of furcular
facet 7,3 X 6,0 mm, depth through sternal facets 8,6 mm; humerus— greatest

EARLY PLIOCENE MARINE AVIFAUNA 159

diagonal diameter of brachial depression 14,2 mm, length of dorsal condyle
8,7 mm.

These specimens differ from Morus and resemble Sula in the following
characters: much more expanded ventral lip and smaller dorsal lip of the sternal
facet of the coracoid; more rounded rather than ovoid furcular facet; dorsal
condyle of humerus not noticeably hooked. Compared to the modern species of
Sula, the South African species is small, falling within the lower part of the size
range of the living species Sula sula but slightly exceeding in size the smallest
individuals of that species from the Central Pacific. The material is too
fragmentary for detailed comparisons with living species and although the fossil
form was fairly similar to Sula sula it differs in having the dorsal lip of the sternal
facet of the coracoid narrower and the brachial depression of the humerus
shallower.

It has been suggested that this species of Sula may have been an incidental
warm-water element in the early Pliocene fauna of the south-western Cape, for
which there is precedent among molluscs as well (Olson 1983). The only sulid in
the Cape region today is the endemic gannet Morus capensis, a much larger form
for which no antecedent has been found in the Langebaanweg or Duinefontein
deposits. The modern species of Morus are found in the cool-temperate waters of
the North Atlantic, South Africa, Australia and New Zealand. The genus also
persisted into the late Pleistocene in the North Pacific but died out there
subsequently. In the Miocene and Pliocene of the western North Atlantic, the
sulid fauna consisted mainly, if not entirely, of a variety of species of Morus
differing greatly in size, suggesting that the species of Sula were mainly of tropical
distribution at that time, as they are today. Thus, I would postulate that Morus
probably did not disperse to the Southern Hemisphere until after the early
Pliocene. The discontinuous distribution of the three living species reflects the
discontinuity of suitable habitat. Dispersal between these widely disjunct
breeding ranges appears to present few problems for these strong-flying birds, as
documented by the numerous instances of vagrant individuals of one species
being found in breeding colonies of another (Crawford et al. 1983).

Family Phalacrocoracidae

Fossil cormorants from Langebaanweg are treated by James (in prep.),
whose identifications are followed here.

Phalacrocorax sp., medium-sized

Material

_Cranial end of right scapula, MBD540; scapular end of right coracoid,
MBD242; part of humeral end of right coracoid, MBD252; shafts of right
coracoids, MBD382, MBD567; shaft and sternal end of left coracoid, MBD383;
proximal end of right humerus, MBD326; proximal ends of right ulnae, MBD249,

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

MBD412; proximal end of left ulna, MBD247; distal ends of left ulnae, MBD338,
MBD403; proximal end of left radius, MBD404; proximal end of left carpometa-
carpus, MBD327; shaft and distal end of right carpometacarpus, MBD196; distal
end of left carpometacarpus, MBD329; proximal end and shaft of left femur,
MBD243; proximal ends of left femora, MBD145, MBD248; proximal end of
right tarsometatarsus, MBD417. Less diagnostic specimens that probably also
belong to this species: part of humeral end of left coracoid, MBD385 (small);
shaft of right humerus, MBD333A; abraded distal end of left tarsometatarsus,
MBD235 (small). Minimum number of individuals, 3.

Remarks

This material is considered to be conspecific with the species from
Langebaanweg described by James (in prep.), although a few very minor points
of osteological difference do exist between the Duinefontein and Langebaanweg
_samples. James (in prep.) discusses the evolutionary relationships of this fossil
species.

The amount of size variation in bones of medium-sized cormorants from
Duinefontein is comparable to that observed in the much larger sample of fossil
cormorants from Langebaanweg. The proportion of larger and smaller indi-
viduals of this species differs among the major stratigraphic members of the
Varswater Formation at Langebaanweg, suggesting that a larger and a smaller
population of the species may have coexisted on the South African coast during
the early Pliocene (James in prep.). If this view is correct, then it is apparent that
individuals from both populations were deposited at Duinefontein as well as at
Langebaanweg.

Phalacrocorax cf. (Microcarbo) sp.

Material
Distal end of left ulna, MBD245.

Remarks

This ulna is small enough to fall within the size range of Phalacrocorax
(Microcarbo) coronatus, the living endemic marine ‘microcormorant’ of South
African waters. Unfortunately, the specimen is not sufficiently diagnostic to allow
positive identification. An examination of variation in long bones of recent
Phalacrocorax capensis showed that the distal end of the ulna is especially likely
to be atypically small in odd individuals (James in prep.). The chance that this
bone belonged to just such an odd individual of the medium-sized cormorant
cannot be entirely ruled out.

Nevertheless, as the presence of a small cormorant in southern Africa during
the late Pliocene is affirmed by two bones from the Varswater Formation at
Langebaanweg (James in prep.), the ulna from Duinefontein could well belong to
the same species.

EARLY PLIOCENE MARINE AVIFAUNA 161

Order GALLIFORMES
Family Phasianidae

Francolinus sp.

Material
Scapular end of right coracoid, MBD544.

Remarks

The most abundant bird at Langebaanweg is a medium-sized species of
francolin (Rich 1980) about the size of Francolinus africanus. Although the single
specimen from Duinefontein is not diagnostic at the species level within the genus
Francolinus, it is nevertheless identical with coracoids of the abundant francolin
at Langebaanweg and very likely referable to the same species.

DISCUSSION

Previous to the South African discoveries, there was practically nothing
known about sea-birds in the Tertiary of the Southern Hemisphere, apart from
numerous reports of fossil penguins and a few bones of pseudotoothed birds
(Pelagornithidae, Pelecaniformes—see Olson in press). Hence the Duinefontein
fauna, despite the relative paucity of specimens and their poor preservation,
represents a significant addition to our knowledge of Tertiary marine birds.

The species composition at Duinefontein is contrasted with that of marine
birds at Langebaanweg and Ysterplaat in Table 1. The differences between these
sites are probably due almost entirely to the nature of the depositional
environment. That at Ysterplaat was most likely a high-energy beach deposit, as
only the very dense, durable bones of the three larger species of penguins were
found, and these are heavily abraded. At Duinefontein, conditions for preserva-
tion were somewhat better, probably reflecting the alternation between beach
and lagoon postulated by Rogers (1979).

The five species found at Duinefontein that are absent at Langebaanweg
(Oceanites sp., Procellaria sp., Calonectris sp., Puffinus spp. A and C) are all
likely to have been non-breeding migrants that died at sea and washed ashore.
Representatives of each of these genera occur regularly in Cape waters today
(Brooke 1981). Oceanites oceanicus and Procellaria aequinoctialis, which may be
closest to the species of Oceanites and Procellaria at Duinefontein, occur in
Antarctic and cold Subantarctic waters during summer, but move northward to
Subtropical waters of high productivity, including those off South Africa, in the
southern winter. Calonectris diomedea, on the other hand, breeds in Subtropical
waters of the North Atlantic and occurs regularly in Subtropical waters off South
Africa in the southern summer. If such patterns of distribution had been
established by the early Pliocene, it might indicate that deposition at Duinefon-
tein was not restricted to a particular season of the year.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Distribution of fossil Sphenisciformes, Procellariiformes, and Pelecaniformes in the south-

western Cape Province (based on the present paper; Olson 1985, in prep.; James in prep.).

YS = Ysterplaat, DF=Duinefontein, LG=Langebaanweg, GM=Gravel Member,

QSM = Quartzose Sand Member, PPM = Pelletal Phosphorite Member (Beds 3aN and 3aS).
See Hendey (19816) for terminology of the Langebaanweg sequence.

LG LG
LG IEG: PPM PPM
GM QSM 3aS 3aN

eo)
ss)

Species YS

‘Nucleornis’ insolitus x
‘Dege’ hendeyi x
‘? Palaeospheniscus’ huxleyorum x
‘Inguza’ predemersus —
Diomedea sp. ==
Oceanites sp. —
Oceanites zaloscarthmus —
- Fulmarinae, gen. et sp. indet. —
Procellaria sp. —
Calonectris sp. —
Pachyptila salax —
Pachyptila species B —
Pachyptila species C —
Puffinus species A —
Puffinus species B —
Puffinus species C —
Pelecanoides cymatotrypetes —
Sula sp. —
Phalacrocorax medium sp. —
Phalacrocorax small sp. —

[sexe 5|

|
bse bh ex x Se |
| <|

Xx lex x X XxX XK kK Xx | Oe x
| |

ee SG | cll SKE X= |
ee lexi |
x le Fel) s| e RA xexex

| xx]
x

Puffinus spp. B and C might be analogous to Puffinus griseus and P. puffinus,
each of which occurs as a migrant in Cape waters today. These modern species
breed in the Southern and Northern hemispheres respectively. Both of the
modern species are somewhat more specialized in wing morphology than the
Pliocene forms, whose similarities to Puffinus tenuirostris may be due to shared
primitive characters and may not necessarily indicate close relationship with that
strictly Pacific species. Puffinus sp. A seems not to have any close living relatives
and may represent an extinct lineage, the oceanographic preferences of which
could not then be inferred. Not much can be said about the enigmatic fulmarine
except that in the Southern Hemisphere the fulmarines breed only in Antarctic or
Subantarctic waters. Of possible relatives, Daption and Fulmarus occur regularly
off the Cape today.

Three species of marine birds are found only at Langebaanweg and are
absent at Duinefontein. However, because Diomedea sp. and Pelecanoides
cymatotrypetes are known only from one and three bones, respectively, their
absence from Duinefontein is probably attributable to chance alone. It is not at all
clear why albatrosses should be so scarce or absent in these deposits, considering

EARLY PLIOCENE MARINE AVIFAUNA 163

their relative abundance in Neogene marine deposits in the Northern Hemisphere
and given the abundance of albatrosses in South African waters today.

Another matter is the complete absence of the storm-petrel Oceanites
zaloscarthmus at Duinefontein, whereas at Langebaanweg it is the second most
abundant procellariiform bird (Olson 1985). Only three other fossils of
Oceanitidae have so far been reported from Tertiary marine deposits anywhere
(Olson 1985, in press), so the Langebaanweg collections have increased the total
world sample by a factor of nearly 60. The very small size, the highly pelagic
nature of their existence, and the non-diving habits of storm-petrels probably
contribute to their scarcity as fossils. For so many remains to be recovered from
Langebaanweg argues for exceptional circumstances of fossilization. In this case,
these storm-petrels probably died in the quiet waters of an estuary in the vicinity
of a breeding island (Olson 1983, 1985), circumstances that would have reduced
the likelihood of deposition of pelagic, offshore migrant species, thus accounting
in part for the differences in species composition observed between Duinefontein
and Langebaanweg.

ACKNOWLEDGEMENTS

At the South African Museum I was greatly assisted in all aspects of my
research by Philippa Haarhoff and Q. Brett Hendey. My stay in Cape Town was
made possible through the Percy FitzPatrick Institute of African Ornithology,
University of Cape Town, through Timothy M. Crowe. Funding for my research
came from the FitzPatrick Institute, the University of Cape Town, the Council for
Scientific and Industrial Research, Pretoria, and the Smithsonian Institution.
Helen F. James supplied information on the cormorants and commented on the
manuscript. Clive Booth of the South African Museum kindly supplied the
photographs. I am indebted to David G. Ainley, Richard K. Brooke, Kenneth
E. Campbell, David C. Duffy, Philippa Haarhoff, Hildegarde Howard, and
David W. Steadman for critical comments on the manuscript.

REFERENCES

Brooke, R. K. 1981. The place of South Africa in the world of seabirds and other marine
animals: 135-147. In: Cooper, J. ed. Proceedings of the Symposium on Birds of the Sea and
Shore. Cape Town: African Sea-bird Group.

CRAWFORD, R. J. M., SHELTON, P. A., Cooper, J. & Brooke, R. K. 1983. Distribution,
population size and conservation of the Cape gannet Morus capensis. S. Afr. J. mar. Sci. 1:
153-174.

DINGLE, R. V., Lorp, A. R. & HENDEY, QO. B. 1979. New sections in the Varswater Formation
(Neogene) of Langebaan Road, south-western Cape, South Africa. Ann. S. Afr. Mus. 78:
81-92.

DINGLE, R. V., SIESSER, W. G. & Newton, A. R. 1983. Mesozoic and Tertiary geology of
southern Africa. Rotterdam: Balkema.

HENDEY, Q. B. 1981la. Geological succession at Langebaanweg, Cape Province, and global
events of the late Tertiary. S. Afr. J. Sci. 77: 33-38.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

HENpDEY, QO. B. 1981b. Palaeoecology of the late Tertiary fossil occurrences in ‘E’ Quarry,
Langebaanweg, South Africa, and a reinterpretation of their geological context. Ann. S.
Afr. Mus. 84: 1-104.

Henpey, Q. B. 1982. Langebaanweg. A record of past life. Cape Town: South African Museum.

JAMES, H. F. In prep. Fossil cormorants (Aves: Phalacrocoracidae) from Cape Province, South
Africa.

Muizon, C. pe. 1980. Des baleines dans le désert! In: Muséum National d Histoire Naturelle:
Récits et Découvertes: 183-190. Paris: Fernand Nathan.

Muizon, C. DE. 1981. Les vertébrés fossiles de la Formation Pisco (Pérou). Premiére partie:
Deux nouveaux Monachinae (Phocidae, Mammalia) du Pliocéne du Sud-Sacaco. Trav. Inst.
fr. Etud. andines 22: 1-161. In: Recherches sur les grandes Civilisations, Mém. N° 6. Paris:
A.D.P.E.

Oxson, S. L. 1983. Fossil seabirds and changing marine environments in the late Tertiary of
South Africa. S. Afr. J. Sci. 79: 399-402.

Otson, S. L. 1985. Early Pliocene Procellariiformes (Aves) from Langebaanweg, south-western
Cape Province, South Africa. Ann. S. Afr. Mus. 95: 123-145.

Otson, S. L. In press. The fossil record of birds. Jn: FARNER, D., Kinc, J. & PARKES, K. C. eds.
Avian biology. Vol. 8. New York: Academic Press.

Otson, S. L. In prep. A revision of South African fossil penguins (Aves: Spheniscidae).

_ Ricu, P. V. 1980. Preliminary report on the fossil avian remains from late Tertiary sediments at
Langebaanweg (Cape Province), South Africa. S. Afr. J. Sci. 76: 166-170.

Rocers, J. 1979. The sedimentary succession at the Koeberg nuclear power station,
Melkbosstrand. Abstr. 18th Congr. geol. Soc. S. Afr. 1: 310-322.

SrESSER, W. G. 1980. Late Miocene origin of the Benguela upswelling [sic] system off northern
Namibia. Science 208: 283-285.

Simpson, G. G. 1971. Fossil penguin from the late Cenozoic of South Africa. Science 171:
1144-1145.

Simpson, G. G. 1973. Tertiary penguins (Sphenisciformes, Spheniscidae) from Ysterplaats, Cape
Town, South Africa. S. Afr. J. Sci. 69: 342-344.

Simpson, G. G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene of
Langebaanweg, Cape Province, South Africa. Ann. S. Afr. Mus. 69: 59-72.

Simpson, G. G. 1979a. A new genus of late Tertiary penguin from Langebaanweg, South Africa.
Ann. S. Afr. Mus. 78: 1-9.

Simpson, G. G. 1979b. Tertiary penguins from the Duinefontein site, Cape Province, South
Africa. Ann. S. Afr. Mus. 79: 1-7.

TANKARD, A. J. 1975a. The marine Neogene Saldanha Formation. Trans. geol. Soc. S. Afr. 78:
257-264.

TANKARD, A. J. 1975b. The late Cenozoic history and palaeoenvironments of the coast margin of
the south-western Cape Province, South Africa. Rhodes University: Unpublished Ph.D.
thesis.

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
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STORRS L. OLSON

AN EARLY PLIOCENE MARINE AVIFAUNA
FROM DUINEFONTEIN,
CAPE PROVINCE, SOUTH AFRICA

95 PART 5 JUNE 1985 | ISSN 0303-2515

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For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).

Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FISCHER, P.-H., DUvAL, M. & RaFFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ENNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
June 1985 Junie
Part 5 Deel

CRETACEOUS FAUNAS FROM
AVEUVLAND AND NATAL, SOUTH AFRICA
THE AMMONITE FAMILY
ROsSsMATICERATIDAE SPATE, 1922

By

WILLIAM JAMES KENNEDY
&
BERBERT CHRISTIAN KLINGER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
THE AMMONITE FAMILY KOSSMATICERATIDAE SPATH, 1922

By
WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford
&

HERBERT CHRISTIAN KLINGER

South African Museum, Cape Town

(With 34 figures)

[MS accepted 27 June 1984]

ABSTRACT

The occurrence of members of the family Kossmaticeratidae in South Africa is well known,
and belies the rarity of the group.

The following are described below: Marshallites cf. cumshewaensis (Whiteaves, 1884), which
is new to South Africa and only the second record of the genus in the Southern Hemisphere;
Kossmaticeras (Kossmaticeras) theobaldianum (Stoliczka, 1865) and varieties, K. (K.) sparsicos-
tatum (Kossmat, 1897), K. (K.) sakondryense Collignon, 1954, K. (K.) jonesi Collignon, 1965,
K. (K.) jeletzkyi Collignon, 1965, K. (Natalites) africanus (van Hoepen, 1920) of which K. (N.)
natalensis (Spath, 1922) is a synonym, K. (N.) faku (van Hoepen, 1920) of which K. (N.)
acuticostatus (Spath, 1922) is a synonym, K. (N.) similis Spath, 1921, K. (N.) elegans sp. nov.,
K. (Karapadites) karapadensis (Kossmat, 1897), K. (K.) cf. madrasinus (Stoliczka, 1865), K. (K.)
besairiei Collignon, 1954, K. (K.) planissimus Collignon, 1966, Maorites cf. subtilistriatus
Collignon, 1954 (the first record of the genus from South Africa), Gunnarites antarcticus (Weller,
1903) and G. kalika (Stoliczka, 1865).

CONTENTS

PAGE
LiSAAROCGRTICHTCON Vaeee ter ae er ttee & cig nee Ce ae mt ee Sere aR aL ere Rr eee 165
OCAN OMOPS PCIE Se cine) hos eg ee else bie. ha wien Mere ee 166
te REO CAlNLIC Siycpey rae Rete Pes adn cat eC de a Son eee Pe hee 166
SEAT OCY D SokS e e ue a ne a a a ae a ie a 166
DIMENSIONS OLSPECIMENS 26 2 adoe-h ele ace tee ak i Mean: 167
SULUNE Te MMMM OLO CWA etre ces ce ts a a ake eee, Soest choo eper a 167
Systematic palacontology 225.05 er see oe 2s Gee ne Soe 167
FR CKMOWIC CS CMENES eh sare sc ee mine mu ae a Me es Loe ar eee 229
INGKEREN CES ie cated ce otise cu gona etann SRE ot Oe vege 230

INTRODUCTION

The Kossmaticeratidae are a highly distinctive family of Desmocerataceae
that have their probable origins in the late Aptian and range to the Maastrichtian.

165

Ann. S. Afr. Mus. 95 (5), 1985: 165-231, 34 figs.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

The distinctive features of the group are the ornament of fine to coarse, often
dense ribs (sometimes associated with tubercles), which are interrupted and often
truncated by oblique constrictions. The group is best known from around the
Indian and Pacific oceans, especially in southern India, Madagascar, Japan, and
New Zealand, although it ranges widely. Its presence in South Africa has been
well known since the publications of Woods (1906), Van Hoepen (1920, 1921)
and Spath (1921a, 1921b, 1922) but in contrast to other groups described by the
present authors, the previously published records give an unbalanced view of its
occurrence, for it 1s rare.

Extensive reviews of the Kossmaticeratidae are given by Collignon (1954,
1955), where all species described up to that date are listed. Important new faunas
are described by Collignon in the Atlas (1964, 1965a, 1965b, 1966, 1969, 1970,
1971) and by Henderson (1970). In view of the scarcity of kossmaticeratids in
South Africa, which adds nothing to our knowledge of the evolution of the
group—although their presence clarifies stratigraphic and geographic distribu-
tions—no general discussion is provided below.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of the
material studied:

BMNH British Museum (Natural History)

DM Durban Museum

GSC Geological Survey, Canada, Ottawa

NMB_ National Museum, Bloemfontein (on permanent loan to SAM)
SAM South African Museum, Cape Town

SAS South African Geological Survey, Pretoria

TM Transvaal Museum

UD University of Natal, Durban; Geology Department Collection.
YPM ~~ Peabody Museum, Yale University.

FIELD LOCALITIES

Details of localities mentioned in the text are given by Kennedy & Klinger
(1975); fuller descriptions of sections are deposited in the Palaeontology
Department of the British Museum (Natural History), London; Geological
Survey, Pretoria; and South African Museum, Cape Town.

STRATIGRAPHY

Kennedy & Klinger (1975) proposed a series of working divisions of the
Barremian to Maastrichtian of Zululand, deferring erection of a detailed
biozonation until revision of the ammonite faunas was completed. At the same
time they admitted that the stage divisions recognized were ‘local’ only, because
of problems of interpretation of these stages in the type areas of western Europe

CRETACEOUS FAUNAS FROM SOUTH AFRICA 167

and correlation from the type areas to southern Africa. Recent work has shown
that, in the case of the Campanian—Maastrichtian boundary, the limit has been
drawn too high in the sequence. Division Campanian IV, from which Saghalinites
cala (Forbes), Pachydiscus (Pachydiscus), Gunnarites antarcticus (Weller),
Nostoceras sp. and Pachydiscus (Neodesmoceras) were recorded, is Lower
Maastrichtian. The Pachydiscus (Pachydiscus) is in fact P. (P.) neubergicus
(Hauer), an exclusively Maastrichtian species, while the Pachydiscus (Neodes-
moceras) 1s P. (N.) mokotibensis Collignon, also exclusively Maastrichtian. The
succeeding Campanian V is thus also Maastrichtian, as is confirmed by the
presence of a specimen of Eubaculites latecarinatus (Brunnschweiler) at this
horizon at locality 118. The authors will continue to use the existing scheme
modified to:

Maastrichtian a (= ‘Campanian’ IV) and Maastrichtian b (= ‘Campanian’ V).

DIMENSIONS OF SPECIMENS

All dimensions given below are in millimetres:
D = diameter, Wb = whorl! breadth, Wh = whorl height, U = umbilical diameter;
c and ic refer to costal and intercostal measurements respectively.

Figures in parentheses are dimensions as a percentage of the total diameter.

SUTURE -EERMINOEOGY

The suture terminology of Wedekind (1916), reviewed by Kullmann &
Wiedmann (1970) is followed here:
I = internal lobe, U = umbilical lobe, L = lateral lobe, E = external lobe.

SYSTEMATIC PALAEONTOLOGY
Superfamily DESMOCERATACEAE Zittel, 1895
Family Kossmaticeratidae Spath, 1922
Subfamily Marshallitinae Matsumoto, 1955
Genus Marshallites Matsumoto, 1955

Type species

Marshallites compressus Matsumoto, 1955, by original designation.

Marshallites cf. cumshewaensis (Whiteaves, 1884)
Fig. 1B-—C

Compare

Haploceras cumshewaensis Whiteaves, 1884: 208, pl. 24 (fig. 1).

Holcodiscoides cumshewaensis (Whiteaves): Imlay & Reeside, 1954: 230.

Marshallites cumshewaensis (Whiteaves): Matsumoto, 1959: 63, pl. 17 (figs 1-4),
ple 19 Gie- 2), pl. 20 (ig: 2); text-iig. 10. McLearn, 1972: 53, pl. 3 (fies 1—2).

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

By monotypy: GSC 4973, from the north shore of Cumshewa Inlet, British
Columbia, Canada.

Material

SAS Z1088, from locality 145, degraded bluffs on eastern side of the
Msunduzi River, Zululand, St. Lucia Formation, Coniacian II.

Description

The specimen retains part of the body chamber and some recrystallized shell
material; the maximum preserved diameter is 39,9 mm. Coiling is moderately
involute; the whorls are compressed, with the greatest breadth just below mid-
flank. The sides are gently inflated, and converge to abruptly rounded
ventrolateral shoulders and a distinctly flattened venter.

Ornament consists of abundant fine, dense prorsiradiate ribs that arise in
bunches from weak umbilical bullae. They sweep forward and are straight across
the inner flank, flex gently backward at mid-flank and are convex, sweeping
forward over the ventrolateral shoulder to cross the venter in a broad convexity.
They branch at or about mid-flank, and there are occasional intercalated short
ribs so that there are many more ribs than umbilical bullae. There are numerous
flexuous, prorsiradiate constrictions, associated with adapical and adapertural
collar ribs that are slightly stronger than the remaining ribs and oblique to the ribs
behind them.

The sutures are not exposed.

Discussion

The specimen closely resembles specimens of Marshallites cumshewaensis
figured by Matsumoto (1959). Of other species referred to this genus, M. com-
pressus Matsumoto (1955: 123, pl. 8 (figs 1-2), text-figs 1-2) is more evolute
and compressed, while M. compressus puzosioides Matsumoto (1955: 125, pl. 8
(figs 3—4)) has extremely fine ornament. Marshallites olcostephanoides Matsu-
moto (1955: 129, pl. 8 (figs 5-7), text-fig. 4) is, as the name suggests,
Olcostephanus-like, evolute and with a whorl breadth to height ratio of between
0,85 and 1,2; the flanks rounded and merging with the venter rather flattened with
a distinct ventrolateral shoulder.

Marshallites columbianus McLearn (1972: 54, pl. 3 (fig. 3)) has much coarser
ribs with a more robust whorl. Marshallites papillatus (Stoliczka) (1865: 159,
pl. 77 (figs 7-8)) is more evolute, with a less compressed whorl, coarser ribs and
many strong constrictions that are far more prominent than in our species.

Occurrence

Marshallites cumshewaensis is an Albian to Cenomanian species, best known
from British Columbia and Alaska. It has not been previously recorded in the

169

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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sISUaDMAYSUIND “Jd San|oOYSAV ‘Q- ‘purjnynz ‘soy Awpeoo] woy ¢/VOZIH SVS ‘S96I ‘UOUSIT[OD snuissiunjd (sanpodvivy) SpdoaoUssoy

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170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Southern Hemisphere, although the genus may occur in New Zealand (Hender-
son 1970).

Subfamily Kossmaticeratinae Spath, 1922
Genus Kossmaticeras de Grossouvre, 1901

Subgenus Kossmaticeras de Grossouvre, 1901

Type species
Ammonites theobaldianus Stoliczka, 1865, by original designation of De
Grossouvre (1901).

Kossmaticeras (Kossmaticeras) theobaldianum theobaldianum Stoliczka, 1865
Figs 1D-E, 2A-E
-Kossmaticeras theobaldi Stoliczka: Collignon, 1955: 20, pl. 1 (figs 2-3), pl. 2 (fig. 1) (with

synonymy); 1965b: 24, pl. 423 (figs 1753-1755).
Kossmaticeras theobaldianum (Kossmat): Sastry, Rao & Mamgain, 1968: pl. 4 (figs 1-2).

Material

NMB D943, SAM-—4909 (the original of Kossmaticeras (Madrasites) bhavani
Spath (non Stoliczka), 1921a: 299, pl. 24 (fig. 8)), UD 45A-B, St. Lucia
Formation, Coniacian I, the Skoenberg region, Zululand. BMNH C83329 from
locality 63, also on the Skoenberg, St. Lucia Formation, Coniacian I.

Dimensions

D Wb Wh Wb:Wh U
SAM-—4909 64,1 22, 3(84-7), 23;2 (Gori) 0,96 22,0(34,3)
SAS D943 60,0 VIM BOI) ZRO(SS.0) 1,03 Zi O(G520)
Description

All the specimens available retain variably corroded, recrystallized shell; all
are septate throughout.

The coiling is evolute (less than half the previous whorl is covered) and the
whorls expand slowly. The umbilicus is of moderate breadth (around 35 per cent
of the diameter) and depth, with a flattened wall at approximately 90 degrees to
the flanks of the preceding whorl. The umbilical shoulder is narrowly rounded,
the whorl section slightly, if at all, compressed (whorl breadth to height ratio
varies between 1,03 and 0,96), with somewhat flattened, rounded convergent
flanks and a broadly rounded venter, the greatest breadth being at or a little
outside the umbilical shoulder. SAM-—4909 has approximately 40 primary ribs per
whorl at a diameter of 40 mm. The ribs arise singly or in pairs on the umbilical
shoulder where they are narrow and sharp, with occasional incipient bullae. They
are narrower than the interspaces, prorsiradiate and straight to gently flexed
across the flanks, crossing the venter in a broad shallow convexity. They either

CRETACEOUS FAUNAS FROM SOUTH AFRICA al

Cc. — | D E

Fig. 2. A-—E. Kossmaticeras (Kossmaticeras) theobaldianum theobaldianum (Stoliczka, 1865).

A-B, D-E. UD 45A-B (ex M. R. Cooper Coll.), from the Coniacian of the Skoenberg,

Zululand. C. SAM-—4909, the original of Spath (1921a: 299, pl. 24 (fig. 8)), from the same area.
All x 1.

WZ ANNALS OF THE SOUTH AFRICAN MUSEUM

branch at various points on the flank or are accompanied by shorter intercalated
ribs that also arise at various points on the flank, so that there are approximately
twice as many ribs per whorl over the venter as there are at the umbilical
shoulder. There are periodic constrictions, four or five per whorl; narrow and
deep, they are flanked by strengthened collar-ribs which usually branch twice.

Discussion

Evolute coiling and dense, wire-like ribbing characterize this species, and the
specimens discussed here closely recall the Indian type material. A number of
varieties have been attached to this species: Kossmaticeras (Kossmaticeras)
theobaldianum var. crassicostata Collignon, 1954, discussed fully below, differs
from typical forms in having fewer and more distant, coarse ribs, and is clearly no
more than a variant. In contrast Kossmaticeras theobaldianum paucicostatum
Matsumoto, 1955 (p. 147, pl. 9 (figs 1-2)), a paratype of which is illustrated here
‘as Figure SA—B, has very distant, broad ribs and a rather massive whorl, and
recalls the K. (K.) sparsicostatum (Kossmat, 1897)—K. (K.) pachystoma
(Kossmat, 1897) group, the coarse ribbing of all of which distinguish them from
K. (K.) theobaldianum theobaldianum. Kossmaticeras (K.) japonicum Matsu-
moto, 1955 (p. 150, pl. 9 (fig. 3)), the holotype of which is reillustrated here as
Figure 5C—F, has distinctive low, broad, crowded ribs quite unlike the wiry
ribbing of the present form. Kossmaticeras (K.) recurrens (Kossmat, 1897) (p. 37
(144), pl. 7 (18) (figs 2-3)) has numerous fine ribs, arising at the umbilicus
without bullae, dichotomously branched on the flanks and crossing the venter
with a marked forward projection; there are five constrictions per whorl.
Kossmaticeras (K.) manasoaense Collignon, 1954 (p. 22, pl. 5 (fig. 1)) (see
Fig. 11A-B) and K. (K.) sakondryense Collignon, 1954 (p. 22, pl. 5 (figs 2-6)
(see Fig. 8C-—D) are more compressed, narrowly umbilicate and feebly ribbed
species, while K. (K.) pavlowskyi Collignon, 1954 (p. 24, pl. 2 (figs 2—3)) (see
Fig. 8A—B) is higher-whorled with fine flexuous ribs that are bi- and triplicate at
mid-flank, so that there are three times as many ribs on the venter as at the
umbilicus. Kossmaticeras (K.) virgatitiforme Collignon, 1965b (p. 27, pl. 425
(fig. 1763)) is characterized by a distinctive division of ribs into bundles of three,
with additional intercalatories. Kossmaticeras (K.) jonesi Collignon, 1965b (p. 29,
pl. 426 (figs 1764-1765)) is a flat-sided species, rather bluntly ribbed, and is,
according to Collignon, especially characterized by shallow, progressively
widening constrictions, four per whorl. Kossmaticeras (K.) jeletzkyi Collignon,
1965b (p. 29, pl. 426 (fig. 1766)) was particularly characterized by the presence of
seven constrictions per whorl.

Occurrence

Kossmaticeras (K.) theobaldianum theobaldianum is restricted to the lowest
division of the Coniacian recognized in Zululand, and occurs only in the
Skoenberg region. It was originally described from the Coniacian of southern
India, and is also recorded from the Middle Coniacian of Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 173

Kossmaticeras (Kossmaticeras) theobaldianum crassicostata Collignon, 1954
Figs 3—4

non Ammonites theobaldianus Stoliczka, 1865: 161 (pars), pl. 78 (fig. 3-3a only).

non Holcodiscus theobaldianus Stoliczka, Grobberippte Varietat: Kossmat, 1897: 36 (143).

Kossmaticeras theobaldi Stoliczka, var. crassicostata Collignon, 1954: 17, pl. 1 (fig. 3), pl. 2
(fig. 1); 1955: 21, pl. 1 (fig. 3), pl. 2 (fig. 1); 19655: 24, pl. 423 (fig. 1756), pl. 424 (fig. 1757).

Type
Holotype by original designation, the specimen figured by Collignon (1954,
pl. 2 (fig. 1)) from the Coniacian of Ampozalaoka, Madagascar.

Material

BMNH C83330, from locality 13, hill slopes below Riverview Compound,
750 m north of the sugar-cane railway bridge across the Mfolozi, south of
Mtubatuba, Zululand, St. Lucia Formation, Coniacian II.

Description

The specimen is a beautifully preserved, wholly septate, fragmentary
individual with an estimated original diameter of 50 mm. It retains well-preserved
recrystallized shell.

Coiling is moderatedly evolute, less than half the previous whorl being
concealed, with a fairly deep umbilicus that comprises an estimated 30 per cent of
the total diameter. The whorl section is equidimensional in section. The greatest
breadth is at the umbilical bulla, the flanks are broadly rounded or flattened,
merging with a more narrowly rounded venter. |

Strong umbilical bullae give rise to single or, more rarely, to pairs of primary
ribs. These are narrow and distant, prorsiradiate, passing straight across the inner
flank, thereafter flexing backward across the remainder of the flank and passing

Fig. 3. Kossmaticeras (Kossmaticeras) theobaldianum crassi-
costata Collignon, 1954; BMNH C83330, from locality 13,
Zululand, Coniacian II. xX 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

174

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[BUISIIO oy) ‘AjaLIeA ay} Jo adAyofoy ay} SpS6T ‘UOUSIT[OD vIDISOIISSD4D WNUDIP]DgoaY] (SDABIIDUSSOY) SDABINDUSSOY “py 31F

CRETACEOUS FAUNAS FROM SOUTH AFRICA rs

straight across the venter. Shorter intercalated ribs, arising on the ventrolateral
shoulder, alternate regularly with the primary ribs.

The constrictions are strongly developed and associated with collar-ribs;
the adapical collar is the stronger, arising from an umbilical bulla and branch-
ing into three in characteristic virgatotome fashion; the adapertural collar is
weaker and simple, and followed by a narrow zone of growth lines. In ventral
view the adapical collar forms a much more narrowly rounded peak than the
other ribs.

The sutures are not exposed.

Discussion

The specimen closely resembles the inner whorls of the holotype from
Madagascar (Fig. 4) but differs from the rather bluntly ribbed specimens from
southern India as illustrated by Stoliczka (1865, pl. 127 (figs 2—3)), which we
prefer to refer to Kossmaticeras (K.) theobaldianum paucicostatum, and regard as
transitional to the K. (K.) sparsicostatum—pachystoma group, regarding K. (K.)
crassicostata, with wiry ribs, as closer to the typical form.

Occurrence

Coniacian II of Zululand, Lower (Collignon 1954) or Middle (Collignon
19656) Coniacian of Madagascar.

Kossmaticeras (Kossmaticeras) aff. theobaldianum crassicostata
Collignon, 1954

Fig. 6F

Compare
Kossmaticeras theobaldi Stoliczka, var. crassicostata Collignon, 1954: 17, pl. 1

(iige:s). pl. 2 (fig. 1).

Material
SAS Z1063, from the Skoenberg area, St. Lucia Formation, Coniacian I.

Discussion

The specimen is a fragment only, septate throughout, with recrystallized and
somewhat corroded test. The style of ornament 1s closely similar to that shown by
Kossmaticeras theobaldianum crassicostata, described above, but the ribs are
Sparser, coarser, with wider interspaces and a greater tendency to branch at or
about mid-flank, with fewer intercalatories and more prominent constrictions.

Occurrence

Coniacian I of Zululand.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kossmaticeras (Kossmaticeras) theobaldianum paucicostatum Matsumoto, 1955
Figs SA-B, 6G—H
Ammonites theobaldianus Stoliczka, 1865: 161 (pars), pl. 78 (fig. 3—3a only).

Holcodiscus theobaldianus Stoliczka, Grobberippte Varietat: Kossmat, 1897: 36 (143).
Kossmaticeras theobaldianum paucicostatum Matsumoto, 1955: 147, pl. 9 (figs 1-2).

Type
The holotype is the original of Matsumoto (1955, pl. 9 (fig. 2)) from the
Coniacian of the Bannosawa, a tributary of the Ikushumbets, Hokkaido, Japan.

Material

SAS Z999, from locality 93, hill slopes on either side of Lots H101-102, ESE
of Hluhluwe, Zululand, St. Lucia Formation, Coniacian II.

- Dimensions
D Wb Wh Wb:Wh U Ribs
SAS Z999 67,5 LISI) ZOU S86) 0,86 PAAO(S 1.11) 60
Description

The specimen is a somewhat worn internal mould retaining traces of shell;
two-thirds of the last whorl is body chamber but it is not clear whether or not the
specimen is adult.

The coiling is moderately evolute, the shallow umbilicus comprising 31 per
cent of the diameter. The low umbilical wall is rounded. There are 21—22 umbili-
cal bullae of variable strength on the outer whorl. These give rise to one or two
strong, prorsiradiate primary ribs, some of which bifurcate, while shorter
intercalated ribs arise on the outer flank, giving a total of 60 per whorl. They pass
straight across the inner flank and are projected forward across the outer flank
and ventrolateral shoulders, crossing the venter with a shallow convexity. There
are six Or seven strong, deep constrictions per whorl. These are prorsiradiate,
passing straight across the flanks and strongly projected over the venter, which
they cross with a narrower convexity than that shown by the ribs. The adapical
collar is bullate, strong, branches in two at the ventrolateral shoulder, the
adapertural branch dividing into two a second time over the venter. The
adapertural collar lacks a bulla, is unbranched, and weaker than the adapical one.

The sutures are not decipherable.

Discussion

A paratype of Kossmaticeras (K.) theobaldianum paucicostatum is shown in
Figure 5A—B, for comparison with the Zululand specimen, which is somewhat
worn. Both have the rather blunt ribbing that, as Matsumoto (1955: 148) noted,
suggests affinity to K. (K.) sparsicostatum. The Zululand specimen closely
resembles Stoliczka’s large specimen (1865, pl. 78 (fig. 3—3a)), especially in the
form of the ribs and the collars associated with the constrictions. It differs from

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CRETACEOUS FAUNAS FROM SOUTH AFRICA

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178 ANNALS OF THE SOUTH AFRICAN MUSEUM

ey

CRETACEOUS FAUNAS FROM SOUTH AFRICA 179

K. (K.) sparsicostatum in having more ribs, and far less striking differentiation
into primaries and secondaries, whereas K. (K.) pachystoma is most easily
distinguished by the inflated shell form and more numerous short ribs.

Occurrence

Coniacian II of Zululand; undifferentiated Coniacian of southern India and
of Japan.

Kossmaticeras (Kossmaticeras) sparsicostatum (Kossmat, 1897)
Fig. 7A—-E

Ammonites denisonianus Stoliczka, 1865: 133 (pars), pl. 66 (fig. 1 only).

Holcodiscus sparsicostatus Kossmat, 1897: 38 (145), pl. 6 (17) (fig. 5).

Kossmaticeras sparsicostatum Kossmat: Collignon, 1954: 19, pl. 3 (fig. 1), pl. 4 (fig. 1); 1955: 22,
pl. 3 (fig. 1), pl. 4 (fig. 1); 1965: 26, pl. 174 (fig. 1758).

Material

SAM-—PCZ6395 figured here as Figure 7A—E, presumed to be from locality
72, degraded river cliffs on the Mzinene River, NNE of Hluhluwe, Zululand,
St. Lucia Formation, Coniacian III.

Dimensions
D Wb Wh Wb: Wh U
SAM-—PCZ6395 82,0 Die Ip 8) 30,6(37,3) 0,91 28,2(34,4)
at 44.5 14,2(31,9) 17,7(69,8) 0,80 13,7 (C058)
Description

The specimen is a well-preserved internal mould; all but the last quarter
whorl is body chamber.

At a diameter of 44,5 mm (Fig. 7A—C) the coiling is moderately involute,
39 per cent of the previous whorl being covered. The umbilicus comprises
30,8 per cent of the diameter and is relatively shallow, with a rounded wall,
undercut on the mould. The whorl section is compressed (whorl breadth to height
ratio is 0,80) with the greatest breadth at the umbilical bulla. The flanks are
flattened and converge slightly to an evenly rounded venter.

There are 20 umbilical bullae of variable strength per whorl, arising as
swellings on the umbilical wall. These give rise to single primary ribs that are
narrow, prorsiradiate, pass straight across the inner flank, curve backward at
mid-flank, where they commonly bifurcate, sweep further back and then forward

Fig. 6 (facing page). A—C. Kossmaticeras (Kossmaticeras) sp. cf. jonesi Collignon, 1965; SAS
Z1587, from locality 92, Zululand, Coniacian Il. D-E. Kossmaticeras (Kossmaticeras) jonesi
Collignon, 1965; SAS H146/7, from locality 13, Zululand, Coniacian II]. F. Kossmaticeras
(Kossmaticeras) aff. theobaldianum crassicostata Collignon, 1954; SAS Z1063, from the
Skoenberg area, Zululand, Coniacian I. G-—H. Kossmaticeras (Kossmaticeras) theobaldianum
paucicostatum Matsumoto, 1955; SAS Z999, from locality 93, Zululand, Coniacian I. All x 1.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. A-—E. Kossmaticeras (Kossmaticeras) sparsicostatum (Kossmat, 1897); SAM-—PCZ6395,
presumably from locality 72, Zululand, Coniacian II. x tee

CRETACEOUS FAUNAS FROM SOUTH AFRICA 181

to cross the venter in a broad convexity. There are also secondary ribs, inserted
low on the flank, giving a total of 65-70 ribs per whorl.

There are seven constrictions per whorl with associated collar-ribs, the
adapical collar branches into two or three, the adapertural one is simple and
slightly weaker.

On the outer whorl, the cross-section is somewhat broader (whorl breadth to
height ratio is 0,91), and the ribbing coarser. Irregular and variably developed
umbilical bullae, eight per half whorl, give rise to a primary rib or a pair of ribs,
and there are also non-bullate primaries. Some primaries bifurcate at mid-flank
and there are also intercalated secondaries, giving a total of 26 ribs per half whorl.
There are four constrictions on the last half whorl of the phragmocone. These are
deep, broad, prorsiradiate and straight on the flanks and projected over the
venter into a narrow convex peak. The adapical collar-rib is strong, arises at a
prominent bulla and branches at mid-flank and on the ventrolateral shoulder. The
adapertural rib is weaker, and simple.

The suture-line is only partially exposed, is deeply and intricately subdivided,
and typical for the genus.

Discussion

The relationship of Kossmaticeras (K.) sparsicostatum and K. (K.) theobal-
dianum theobaldianum, K. (K.) t. crassicostata and K. (K.) t. paucicostatum has
been discussed above. The species is close to K. (K.) pachystoma (Kossmat)
(1897: 39 (146), pl. 7 (18) (fig. 1)), from which it is most easily separated by the
compressed as compared to circular cross-section, narrower and rather irregular
ribbing.

Occurrence

Coniacian of Zululand, Lower (Collignon 1954) or Middle (Collignon 19655)
Coniacian of Madagascar, and undifferentiated Coniacian of southern India.

Kossmaticeras (Kossmaticeras) sakondryense Collignon, 1954
Figs 8C—D, 10A-—B

Kossmaticeras (Kossmaticeras) sakondryense Collignon, 1954: 22, pl. 5 (figs 2-5), (fig. 6 = var.
eboroense); 1955: 22, pl. 5 (figs 2-5), (fig. 6 = var. eboroense): 1965b: 27, pl. 425 (fig. 1761).
Type
The holotype, by original designation, is the original of Collignon (1954, pl. 5

(fig. 5)), reproduced here as Figure 8C—D, from the Coniacian of the Ravin
d’Anjoho, Sakondry Valley, Madagascar.

Material

SAS Z929, from locality 93, hill slopes on either side of boundary between
Lots H101—102, ESE of Hluhluwe. Zululand, St. Lucia Formation, Coniacian II.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

E

Fig. 8. A-B. Kossmaticeras (Kossmaticeras) pavlowskyi Collignon, 1954; a paratype, from the

Coniacian of Ampozalaoka (Menabe), Madagascar, the original of Collignon (1954, pl. 2

(fig. 2-2b)). C-—D. Kossmaticeras (Kossmaticeras) sakondryense Collignon, 1954; the holotype,

the original of Collignon (1954, pl. 5 (fig. 5—5b)), from the Coniacian of Sakondry, Madagascar.

E. Kossmaticeras (Kossmaticeras) sakondryense var. eboroense Collignon, 1954; the holotype of

the variety, the original of Collignon (1954, pl. 5 (fig. 6)), from the Coniacian of Eboro,
Madagascar. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 183

Dimensions

D Wb Wh Wb: Wh U
SAS Z929 12.1 100)= 23-5(32-3)- 2756758) 0,85 21,4(29,4)
Description

The specimen is somewhat abraded, in part an internal mould, in part
retaining well-preserved aragonitic shell material.

Coiling is moderately involute, with 57 per cent of the previous whorl being
covered. The rather shallow umbilicus comprises 29,5 per cent of the diameter.

Fig. 9. Kossmaticeras (Kossmaticeras) jeletzkyi Collignon, 1965; SAS D1342, from locality 72,
Zululand, Coniacian III. x 1.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc D E

Fig. 10. A-B. Kossmaticeras (Kossmaticeras) sakondryense Collignon, 1954; SAS Z929, from
locality 93, Zululand, Coniacian II. C-E. Kossmaticeras (Natalites) elegans sp. nov.; the
holotype, SAS H30/9, from locality 100, Zululand, Santonian I. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 185

The whorl section is compressed (whorl breadth to height ratio is 0,85) with the
greatest breadth close to the narrowly rounded umbilical shoulder. The sides are
flattened, and converge to the rounded ventrolateral shoulders. The venter is
somewhat flattened.

Ornament consists of approximately 100 fine ribs; 30 of these arise at small
umbilical bullae developed from broad swellings on the umbilical wall. The bullae
are fine, comma-shaped, and variable in strength. They give rise to prorsiradiate
ribs that pass straight across the sides, bend forward or are gently flexed on the
inner flank and bend forward across the outer flank and ventrolateral shoulder to
project over the venter in a strong convexity.

These primary ribs branch once or twice on the outer flank and loop across
the venter, while there are also shorter intercalated ribs.

There are nine narrow, deep, prorsiradiate constrictions per whorl. These
are straight on the inner flank but sweep forward over the venter in a deep
convexity on the mould that is scarcely visible when the shell is present.
Strengthened collar-ribs flank the constrictions; the adapical collar subdivides
across the ventrolateral shoulder to give rise to three or four riblets arranged in a
virgatotome pattern.

The sutures are not exposed.

Discussion

Kossmaticeras (K.) sakondryense 1s a distinctive, compressed, high-whorled,
involute and delicately ornamented K. (Kossmaticeras); features which separate
it from most other species of the subgenus. There are some similarities to K. (K.)
Japonicum Matsumoto (1955: 150, pl. 9 (fig. 3)) (see Fig. SC—F), from the
Coniacian of Hokkaido and Saghalien, but the Japanese form is less compressed,
with lower whorls and coarser ribs with fewer (6—7 versus 9) constrictions per
whorl.

Occurrence

Coniacian II of Zululand; Lower (Collignon 1954) or Middle (Collignon
1965b) Coniacian of Madagascar.

Kossmaticeras (Kossmaticeras) aft. sakondryense Collignon, 1954
Fig. 21A—C

Compare

Kossmaticeras (Kossmaticeras) sakondryense Collignon, 1954; herein, p. 181,
Fig. 8C—D.

Material

SAS Z934, from locality 93, hill slopes on either side of boundary of
Lots H101—102, ESE of Hluhluwe, Zululand, St. Lucia Formation, Coniacian II.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions
D Wb Wh Wb:Wh wi
SAS 2934 59,0 UG,5(S1,3)) 25,0423) 0,74 15532559)

Description and discussion

The specimen is a wholly septate internal mould, and somewhat abraded.
The general style of ornament is rather similar to that of Kossmaticeras (K.)
sakondryense, described above, but it has a slightly smaller umbilicus (25,9 per
cent vs 29,4 per cent), for which reason it is separated from the restricted form of
the species.

Occurrence

Coniacian II of Zululand.

Kossmaticeras (Kossmaticeras) jonesi Collignon, 1965
Figs 6D-E, 12C-—D
Kossmaticeras (Kossmaticeras) jonesi Collignon, 1965b: 29, pl. 426 (figs 1764-1765).

Type

The holotype, by original designation, is the original of Collignon (1965),
pl. 426 (fig. 1764)), from the Zone of Kossmaticeras theobaldi and Barroisiceras
onilahyense of Ankinatsy-Souromaraina (Belo-sur-Tsiribihina), Madagascar.

Material

BMNH C83331 and SAS H146/7, from locality 13, hill slopes below
Riverview Compound, 750 m north of the sugar-cane railway bridge across the
Mfolozi, south of Mtubatuba, Zululand, St. Lucia Formation, Coniacian II.

Description

Both the specimens are fragmentary and retain either original aragonitic, or
recrystallized shell material. SAS H146/7 represents an individual with an
estimated adult diameter of 45 mm and a quarter of a whorl of body chamber;
BMNH C83331 is a wholly septate fragment of an individual with an estimated
diameter of 75 mm.

Coiling is moderately evolute, the umbilicus comprising an estimated 30 per
cent of the total diameter, of moderate depth with a flattened wall, sloping
outwards. The whorl section is compressed (whorl breadth to height ratio is
circa 0,85) with the greatest breadth low on the flank or at the umbilical bullae.

Fig. 11 (facing page). A-B. Kossmaticeras (Kossmaticeras) manasoaense Collignon, 1954; the

holotype, the original of Collignon (1954, pl. 5 (fig. 1-1b)), from the Coniacian of Manasoa,

Madagascar. C. Kossmaticeras (Natalites) africanus faku (van Hoepen, 1920); specimen in the

Durban Museum cited by Spath (1921a: 47), from an unspecified horizon in the Umzamba
Formation near the Umzamba Estuary. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 187

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

e

CRETACEOUS FAUNAS FROM SOUTH AFRICA 189

There are numerous closely spaced umbilical bullae, of variable strength,
and these give rise to single or paired ribs, which commonly bifurcate at mid-flank
while shorter intercalatories are also present between primaries. The ribs are
crowded, prorsiradiate and flexuous, convex at mid-flank, concave on the outer
flank and projected across the venter. The constrictions are prominent, relatively
strong, prorsiradiate and flexuous with associated collar-ribs; the adapical one is
strong, arises from an umbilical bulla, and splits into four in typical virgatotome
style; the adapertural one is simple.

The sutures are not exposed.

Discussion

The diagnostic features are the compressed whorl combined with strong,
dense ribbing, the ribs branching on the outer flank, and the prominent strong
constrictions. Together, these readily distinguish the species from the coarsely
ribbed Kossmaticeras (K.) sparsicostatum, inflated K. (K.) pachystoma, delicately
ribbed K. (K.) sakondryense or serpenticone and wiry-ribbed K. (K.) theobal-
dianum group. Kossmaticeras (K.) jeletzkyi Collignon (19656: 29, pl. 426
(fig. 1766)) has not dissimilar proportions, but seven constrictions per whorl, a
marked weakening of ribs at mid-flank, and numerous intercalatories. Kossma-
ticeras (K.) manasoaense and K. (K.) virgatitiforme all differ in having finer ribs,
with individually distinctive styles and branching patterns (see Collignon 1954,
1955, 19656 for details).

Occurrence

Coniacian II of Zululand; Middle Coniacian of Madagascar.

Kossmaticeras (Kossmaticeras) sp. cf. jonesi Collignon, 1965
Fig. 6A—C

Compare

Kossmaticeras (Kossmaticeras) jonesi Collignon, 1965b: 29, pl. 426 (figs 1764-
WO):

Material

SAS Z1587, from the St. Lucia Formation, Coniacian II at locality 92.
Bulldozer scrapings and hill slopes on the farm Panplaas, ESE of Hluhluwe,
Zululand, St. Lucia Formation, Coniacian II or III.

Fig. 12 (facing page). _A-B. Kossmaticeras (Natalites) africanus africanus (van Hoepen, 1920);

the holotype of Madrasites natalensis Spath, 1922, BMNH C19432, from an unspecified horizon

in the Umzamba Formation. C—D. Kossmaticeras (Kossmaticeras) jonesi Collignon, 1965;
BMNH C83331, from locality 13, Zululand, Coniacian II. All x 1.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dimensions

D Wb Wh Wb:Wh Ul
SAS Z1587 SES 12,1(32,4) 14,6(39,1) 0,83 IES SILC)
Description

This small specimen retains iridescent nacreous shell, and is septate to a
diameter of 37,3 mm, with indications of the former presence of more than half a
whorl of body chamber.

Coiling is moderately involute, the umbilicus comprising 31,9 per cent of the
diameter, of moderate depth with a flattened wall and narrowly rounded
shoulder. The whorl section is compressed with a breadth to height radio of 0,83,
the greatest breadth being at the umbilical shoulder, the flanks flattened,
convergent, with an arched venter.

There are approximately 20 variably developed umbilical bullae per whorl.
These give rise to one or two primary ribs that are narrow and rather sharp. They
are prorsiradiate, flexing forward across the inner flank, convex at mid-flank,
thereafter curving backward into a distinct concavity before sweeping forward to
pass almost straight across the venter. Some ribs bifurcate at various positions;
there are also shorter intercalated ribs and occasional non-bullate primaries,
giving a total of approximately 65 ribs per whorl.

There are six constrictions per whorl, marked on the shell by a strong
adapical collar that branches twice, and a series of fine riblets over the site of the
constrictions.

The sutures are not exposed.

Discussion

The holotype of Kossmaticeras (Kossmaticeras) jonesi is much larger than the
present specimen (107 mm vs 37,3 mm), but it shows sufficient similarities to
allow us to tentatively refer the specimen to this species.

Occurrence

St. Lucia Formation, Coniacian II or III of Zululand.

Kossmaticeras (Kossmaticeras) jeletzkyi Collignon, 1965
Fig. 9
Kossmaticeras jeletzkyi Collignon, 19656: 29, pl. 426 (fig. 1766).

Type

The holotype, by original designation, is the original of Collignon (1965),
pl. 426 (fig. 1766)) from the Middle Coniacian of Analabe (Belo-sur-Tsiribihina),
Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 191

Material

SAM-—D1342, from locality 72, degraded cliffs and alluvial flats on north
side of Mzinene River, NNE of Hluhluwe, Zululand, St. Lucia Formation,
Coniacian III.

Dimensions

D Wb Wh Wb: Wh U Ribs
SAM-D1342 OES —(—) 43 ,8(37,6) - 39506325) h 06
Description

The specimen is a somewhat distorted internal mould retaining extensive
areas of recrystallized shell. About two-thirds of the outer whorl are body
chamber.

Coiling is relatively evolute, the shallow, crater-like umbilicus comprising
33,5 per cent of the diameter with a flattened, outward-inclined wall. The
umbilical shoulder is abruptly rounded, the whorl section is compressed (whorl
breadth to height ratio is 0,82), with the greatest breadth at, or just outside, the
umbilical bullae. The inner flanks are flattened and subparallel; the outer
converge to a high arched venter.

The inner whorls are ornamented by 21 somewhat variable umbilical bullae,
which give rise to one, or rarely two, straight prorsiradiate ribs which generally do
not branch in the area of flank exposed.

On the outer whorl, there are 26 umbilical bullae. These give rise to broad,
flexuous prorsiradiate ribs, singly or in pairs. These flex forward across the inner
flank, are convex and flex backward across the mid-flank, where some branch, or
are accompanied by shorter intercalated ribs. All the ribs are concave across the
outer flank, and project forward over the ventrolateral shoulders to cross the
venter in a broad convexity. There are seven constrictions per whorl. These are
strong and deep on the internal mould, but rather less conspicuous where the
shell is present. The adapical collar-rib is the stronger, bullate and bifurcates
three times. The adapertural collar-rib is simple.

The sutures are not exposed.

Discussion

The distinguishing features of this species are the compressed whorl section
and seven prominent constrictions on the outer whorl, thus separating it from the
allied Kossmaticeras (K.) jonesi Collignon.

Occurrence

Lower Coniacian of Madagascar, Coniacian II of Zululand.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Subgenus Natalites Collignon, 1954

Type species
Madrasites natalensis Spath, 1922, by the original designation of Collignon
(1954: 6) (= Holcodiscus africanus van Hoepen, 1920).

Kossmaticeras (Natalites) africanus africanus (van Hoepen, 1920)
Figs 12A-B; 13A-E, G-I; 16G—I; 18B—H
Holcodiscus africanus van Hoepen, 1920: 146, pl. 26 (figs 3-5); 1921: 23.
Holcodiscus africanus? van Hoepen: van Hoepen, 1921: 23.
Madrasites africanus van Hoepen: Spath, 1921a: 48; 1922: 135.
Maarasites natalensis (Crick MS) Spath 1922: 134, pl. 5 (fig. 3).
Kossmaticeras (Natalites) natalensis Spath: Collignon, 1954: 6; 1955: 13; 1966: 8-9, pl. 457
(fig. 1867), pl. 458 (fig. 1868). Wright, 1957: L374, fig. 490 (1).

- Types

The holotype by original designation is TM 578, the specimen figured by Van
Hoepen (1920, pl. 26 (figs 3-5)); paratypes are TM 543-5, all from the Umzamba
Formation ‘at the mouth of the Umzamba River, Pondoland’, precise horizon
unknown. The holotype of Madrasites natalensis is BMNH C19432, the specimen
figured by Spath (1922, pl. 5 (fig. 3)) from an unknown horizon at the Umzamba
River estuary.

Apart from the types, the specimen in the Durban Museum mentioned by
Spath (1921a: 48) (see Fig. 16G—I), SAM-—7105 and 7073, NMB D1697, and SAS
Z1587 and P1416, are all from the same locality as the types.

Dimensions
D Wb Wh Wb:Wh G)

TM 578 (after

Van H.) 44,0 c.12(27,3) c.17(38,6) O71 9 1SSGoNp
SAS P1416 SES 1152(35,2) 11,6C66;4)7 0:97) esiGar)
NMB D1697 59,4 18,0053) 21,0G554) 0586 = 20%sG
BMNH C19432

(after Spath) 81,0 —(34) —(40) 0,75 —(30)
Description

Coiling is moderately evolute (approximately one-third to one-half of the
previous whorl is covered) with a moderately deep umbilicus that varies between

Fig. 13 (facing page). A-E. Kossmaticeras (Natalites) africanus africanus (van Hoepen, 1920).

A-B. BMNH C83335, x 3,3. C. TM 578, the holotype. D. TM 544, paratype. E. TM 544,

paratype. F. Kossmaticeras (Natalites) africanus faku (van Hoepen, 1920); TM 543, holotype.

G-I. Kossmaticeras (Natalites) africanus africanus (van Hoepen, 1920); SAS P1416.

J-K. Kossmaticeras (Natalites) elegans sp. nov.; paratype, the original of Woods (1906, pl. 42,

(fig. 2a—b)). All from an unspecified horizon in the Umzamba Formation at the Umzamba
Estuary. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 193

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

30 and 36 per cent of the diameter. The whorl section is generally compressed,
with the greatest breadth at the umbilical bulla, and with flattened flanks that
converge to a broadly arched venter. There are on average 20 comma-shaped
umbilical bullae per whorl; they arise as broad swellings on the umbilical wall,
and give rise to single ribs or groups of up to four ribs. These are sharp and
narrow, prorsiradiate and flexuous, convex across the inner mid-flank and
concave across the outer, sweeping forward over the venter in a broad convexity.
There are occasional intercalatories, which do not originate in an umbilical bulla.

There are six to seven broad constrictions per whorl, flanked by collar-like
ribs. The adapertural ones are usually simple and without bullae, whereas the
adapical ones show virgatotome style of branching into two or three secondaries.

Ornament on the innermost whorls is generally very weak, with the
constrictions very conspicuous. On the later part of the phragmocone ribbing and
tuberculation become very conspicuous and bold, but weaken again on the body

chamber.

Discussion

It is difficult to satisfactorily separate Kossmaticeras (Natalites) africanus
(van Hoepen) (of which K. (N.) natalensis (Spath, 1922) is a synonym) from
K. (N.) faku (van Hoepen) (of which K. (N.) acuticostatus (Spath, 1922) is a
synonym). Both are poorly represented in terms of numbers, and little accurate
stratigraphic data on their distribution is available. Generally, K. (N.) africanus is
the more compressed, wider umbilicate form with ribs only branching at the
umbilical bullae, except for the virgatotome branching at the constrictions,
whereas K. (N.) faku has a more inflated whorl section, narrower umbilicus and
abundant intercalatory and branching ribs which arise at mid-flank. However,
these extreme forms are connected by numerous transitions (as already
mentioned by Spath (1921a: 47)) so that separation at more than subspecific level
would seem unnecessary. The holotypes of ‘Madrasites natalensis’ Spath
(Fig. 12A—B) and ‘Madrasites acuticostatus’ Spath (Fig. 15A—C) illustrate the
point. ‘Madrasites natalensis’ has coiling similar to Kossmaticeras (N.) africanus
faku, but ornament comparable to that of K. (N.) africanus africanus. ‘Madrasites
acuticostatus’ has coarse ribbing comparable to that of K. (N.) africanus faku, but
it lacks the abundant mid-flank bifurcations and intercalatories, and in this
respect is again closer to K. (N.) africanus africanus.

Occurrence

A single specimen (Klinger & Kennedy 1980) was recovered in situ from
Bed Cll on the southern side of the Umzamba Estuary, and can be dated as
Santonian III. All the other specimens are from unknown horizons in the
Umzamba Formation. In Madagascar the species (as Natalites natalensis) was
recorded from the Lower Santonian, Zone of Texanites oliveti. As yet, the species
is unknown in Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 195

Kossmaticeras (Natalites) africanus faku (van Hoepen, 1920)

Figs 11C, 13F, 14-15, 16A—F

Holcodiscus faku van Hoepen, 1920: 144, pl. 25 (figs 3-4), pl. 26 (figs 1-2).
Madrasites faku van Hoepen: Spath, 1921b: 47; 1922: 135.
Madrasites acuticostatus Spath, 1922: 134, pl. 8 (fig. 2).

Types

The holotype, by original designation, is TM 542, the specimen figured by
Van Hoepen (1920, pl. 25 (figs 3—4)), and the paratype TM 579 (Van Hoepen
1920, pl. 26 (figs 1-2)), both from an unspecified horizon in the Umzamba
Formation ‘near the mouth of the Umzamba River’ (Van Hoepen 1920: 142). The
holotype by monotypy of Madrasites acuticostatus Spath, 1922, from the same
locality and an equally uncertain horizon, is BMNH C19433.

Material

Apart from the types, the two examples from the Umzamba Formation of
Pondoland, housed in the Durban Museum, mentioned by Spath (19215: 47) (see
Fig. 16A—F) were examined.

Dimensions

D Wb Wh Wb:Wh U
Holotype TM 542 202 20,0655) 23,0403) = O87 alos0 sat)
Description

Coiling is moderately involute, covering more than half of the previous
whorls. The whorl section is compressed, with greatest breadth near the umbilical
edge, the flanks converging to a narrower, rounded venter. On the inner whorls
the umbilical wall is nearly vertical, and the umbilical edge well defined. On the
outer whorl the umbilical wall slants outwards and the edge becomes more
rounded.

There are 18 sharp, comma-shaped umbilical bullae on the holotype. From
there arise pairs or trios of sharp, narrow, sinuous ribs, many of which again
bifurcate near mid-flank. Some ribs arise directly on the umbilical edge and follow
a similar course over the flanks. In total there are about 80 ribs per whorl.

Seven distinct constrictions are present on the outer whorl of the holotype.
These are already noticeable on the umbilical wall, and follow a prorsiradiate,
sinuous path over the flanks, with a marked forward flexure over the venter. The
rib adapical of each constriction is thickened, and shows a virgatotome style of
branching into three or four secondaries. On the body chamber ribbing and
umbilical ornament become weaker and more distant.

Discussion

As discussed above, separation of Kossmaticeras (N.) africanus africanus and
K. (N.) africanus faku is difficult in the case of transitional forms such as the

ANNALS OF THE SOUTH AFRICAN MUSEUM

196

‘Tx ‘Alenisq equiezwiy) oy] Je uoNneWIO4
pquiRZWA) oY) UI UOZIIOY poyloodsun ue wo tadAjyesed ‘67S WL ‘(O76 ‘Usdoo0H Ura) nyvf snuvoidf{p (sal]VIDN) SDLdDUSSOY “Q-Y ‘pT “314

2) qo Vv

17

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘| x ‘Alenisq equiezwiy oy) ye UONeUOY equeZWA ay) UL UOZOY payloodsun ue WO ‘TZ6] ‘yIeds snvjsooynov
sauspapow JO adAjojoy oy ‘EeP6l HN ‘(0261 ‘uedeopx uea) nyvf snuvoifo (SA1]DIDN]) SvooNDUSSOY “Q-Y “ST ‘SIA

9 q

ANNALS OF THE SOUTH AFRICAN MUSEUM

198

ES

CRETACEOUS FAUNAS FROM SOUTH AFRICA 199

holotypes of Madrasites acuticostatus Spath, 1922, or Madrasites natalensis Spath,
1922. Typical K. (N.) faku has abundant bifurcations at mid-flank, whereas
typical K. (N.) africanus lacks these.

Occurrence

None of the specimens is precisely localized within the Umzamba Formation
at the Umzamba Estuary, and the subspecies cannot be dated more precisely than
Middle or Upper Santonian to Lower Campanian.

Kossmaticeras (Natalites) similis Spath, 1921

Fig. 17
Madrasites similis Spath, 1921b: 48, pl. 6 (fig. 1).

Type

The holotype by monotypy is the specimen figured by Spath (19215, pl. 6
(fig. 1)) in the collections of the Durban Museum, from an unspecified horizon in
the Umzamba Formation at the Umzamba Estuary.

Material
No additional material of the species is known.

Dimensions
D Wb Wh Wb:Wh U
Holotype (after
Spath) 100 33(33) 38(38) 0,87 34(34)

Description

The holotype lacks the innermost whorls and is preserved as an internal
mould. Coiling is moderately evolute with successive whorls embracing each
other up to about mid-flank. The whorl section is higher than wide, with greatest
width at the umbilical edge, and then tapers slowly to the broadly rounded venter.

Ornament on the phragmocone consists of strong, conical umbilical
tubercles, and ribbing that arises either from the tubercles or intercalates.
Ribbing is weak near the umbilical edge and inner part of the flank, but increases
outwards across the flanks, and is at a maximum across the venter. On the outer
whorl, the ribbing becomes increasingly irregular, the primary ribs arising singly
or in pairs from weak to strong umbilical bullae, with occasional shorter
intercalated ribs. At the greatest diameter preserved, the ribs coarsen and are
blunted.

Fig. 16 (facing page). A-F. Kossmaticeras (Natalites) africanus faku (van Hoepen, 1920); the two

specimens mentioned by Spath (1921: 47) as Madrasites faku van Hoepen. G-I. Kossmaticeras

(Natalites) africanus africanus (van Hoepen, 1920); the specimen mentioned by Spath (1921: 48)

as Madrasites africanus van Hoepen. All from an unspecified horizon in the Umzamba Formation
at the Umzamba Estuary. All x 1.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

oe GO",
eas ss,

Fig. 17. A-—D. Kossmaticeras (Natalites) similis Spath, 1921. B and C are two different ventral
views of the holotype, from an unspecified horizon in the Umzamba Formation at the Umzamba
Estuary. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 201

There are five deep constrictions per whorl, narrow where shell is present,
but broad on the internal mould. They are straight and prorsiradiate across the
inner and middle flank, flex abruptly forward on the outer flank and cross the
venter with a linguoid apertural projection. There are flanking collars; the
adapertural one is narrow and simple, without an umbilical bulla; the adapical
one stronger and commonly split, virgatotome-fashion, into three riblets.

The sutures are not visible.

Discussion

The very distant, narrow strong ribs and large, distant bullae separate this
species readily from all others referred to the subgenus. As Spath (1921b: 48)
noted, Kossmaticeras (N.) similis stands in the same relationship to K. (N). faku
as K. (K.) sparsicostatum does to K. (K.) theobaldianum.

Occurrence

The holotype is from the Umzamba Formation of Pondoland and is
presumably of Middle or Upper Santonian to Lower Campanian age.

Kossmaticeras (Natalites) elegans sp. nov.
Figs 10C—E, 13J-K, 18A

Types

Holotype SAS H30/9, from locality 100, hill slopes alongside track leading
north from Nkundusi, 1,0—1,5 km N of the village, SE of Hluhluwe, Zululand.
St. Lucia Formation, Santonian I. Paratypes are SAM 13100 and 4811 (= Woods
1906, p. 336, pl. 42, (fig. 2a—b)) from an unknown horizon in the Umzamba
Formation of Pondoland.

Dimensions

D Wb Wh Wb: Wh U
SAS H30/9 63.7 18,0(28,3) 23,5(36,8) O77, 15.7293)
Description

The holotype is largely septate and retains much of its nacreous aragonitic
shell.

The coiling is moderately involute, 67 per cent of the previous whorl being
covered. The umbilicus is of moderate breadth (29,3 per cent of the total
diameter) with the umbilical wall sloping outwards, flattened, with an abruptly
rounded shoulder. The whorl section is compressed (whorl breadth to height ratio
is 0,77), with the greatest width at the umbilical bullae. The whorl sides are high,
flattened, and converge to a narrowly rounded venter on the phragmocone that
broadens on the body chamber. Broad ribs arise on the umbilical wall and give
rise to 21 small comma-shaped bullae per whorl. These give rise to bundles of up
to four, and occasionally five, fine prorsiradiate ribs that are straight across the

ANNALS OF THE SOUTH AFRICAN MUSEUM

202

‘Arenisy equiezuic
oY} Je UONPUIOT EQUIBZWIY) 94) Ul UOZTIOY payloodsun ue Woy [TV ‘1 x “S—Eers WL sodAjesed ou} ‘(976, ‘uedaoy ue) snuvoiafo
SNUDIID (SA1]DIDN]) SbADINDUUSSOY “H-G ‘1 X ‘OOLET-IWVS edAjesed ‘‘aou ds suvsaja (saiyvivN) spsvoDUssoy ‘YW ‘81 314

5 4 v

CRETACEOUS FAUNAS FROM SOUTH AFRICA 203

inner flank but sweep backward at mid-flank and thereafter forward to project
strongly over the ventrolateral shoulders, connecting across the venter in a
marked convexity. On the phragmocone there is occasional secondary branching
and intercalation of short ribs, while simple and intercalated ribs become
common on the body chamber, to give a total of 104 ribs per whorl.

There are nine constrictions per whorl, deep on the mould and partially
exfoliated specimens but much less conspicuous where shell is preserved. They
are flexuous and prorsiradiate and flanked by collar-ribs. The adapertural ones
lack umbilical bullae; the adapical ones are low, narrow and show a virgatotome
branching into three.

The sutures are not exposed.

Remarks

The distinct umbilical bullae on the inner whorls, giving rise to groups of ribs,
clearly indicate this to be a species of Kossmaticeras (Natalites). It is the oldest
species so far recorded. It differs from all other described species of the subgenus
in its involution, compression, and high, flat-sided whorls with weak ornament
throughout, all of which separate it from the other South African species. Of the
various New Zealand species described by Henderson (1970), the present form
most closely resembles K. (Natalites) bensoni Henderson (1970: 39, pl. 4 (fig. 3)).
The latter is a larger form, has stronger, persistent umbilical bullae, thin, narrow
ribs that are markedly flexuous, and four constrictions per whorl, rather than the
nine seen in the present species.

Occurrence

St. Lucia Formation, Santonian I, locality 100, Zululand; Umzamba
Formation (precise horizon unknown), Umzamba Estuary, Transkei.

Subgenus Karapadites Collignon, 1954
(= Karapadites Matsumoto, 1955)

Type species
Holcodiscus karapadensis Kossmat, 1897, by original designation of
Collignon (1954).

Kossmaticeras (Karapadites) karapadensis (Kossmat, 1897)
Figs 19C-E, 24A

Holcodiscus karapadensis Kossmat, 1897: 41 (148), pl. 8 (19) (figs 2, 4).
Karapadites karapadensis Kossmat: Collignon, 1954: 27, pl. 6 (figs 1-4); 1955: 27, pl. 6 (figs 1-4);
1969: 69, pl. 541 (fig. 2121).

Types

Kossmat (1897) based this species on two specimens from the Arialoor
Group of Karapady, southern India, in the Warth Collection. The larger
specimen figured by him as plate 8 (19) (fig. 4a—c) is herein designated lectotype.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

BMNH_ C83328 from locality 14, road cuttings below the compound
immediately south of the Msunduzi River, 2,1 km NNE of Mfolozi, south of
Mtubatuba, Zululand, St. Lucia Formation, Campanian I.

Dimensions
D Wb Wh ~ Wb:Wh U
BMNH C83328 33,8 —(—) IZA (Gses)) _ 11363854)
at 28,9 DO(S43) YO(S3.,2)) 1208 LOpS(Z725))
Description

The specimen is a wholly septate internal mould. Coiling is evolute, less than
a third of the previous whorl being covered: The umbilicus comprises 27,5 per
~ cent at a diameter of 28,9 mm, becoming more evolute with growth (33,4 per cent
SEb S)S)ce) 100001).

The whorl section is slightly wider than high with the greatest breadth low on
the flank; the sides are flattened, converging to an arched venter. There are
15 small umbilical bullae per whorl. These give rise to groups of three ribs, almost
invisible on the inner flank, but strengthening across the mid- to outer flank,
where they are joined by intercalated ribs. All are distinctly flexuous and
prorsiradiate, crossing the venter (over which they weaken) in a shallow
convexity. There is a total of 60 ribs per whorl.

Four prominent, strong, deep constrictions are present on the outer whorl.
They are concave and markedly prorsiradiate, crossing the venter with a narrow
linguoid adapertural projection; they weaken over the siphonal line. The
associated adapical collar rib is strong, arises from an umbilical bulla and
bifurcates twice. The adapertural rib is weaker.

The suture-line is shown in Figure 24A, and agrees closely with that of the
lectotype.

Discussion

The single small specimen of Kossmaticeras (Karapadites) karapadensis
agrees well with Kossmat’s type material, and Madagascan specimens illustrated
by Collignon (1954, 1969). The species is distinguished from Kossmaticeras
(Karapadites) madrasinus (Stoliczka, 1865) (p. 139, pl. 70 (figs 1-3)) by the
stronger ribs, well developed on the flank, stronger umbilical bullae, plus six to
eight prominent constrictions per whorl; features differentiating adults are given
by Collignon (1954: 31-32).

Adult Kossmaticeras (Karapadites) besairiei Collignon (1954: 29, pl. 8
(fig. 2); 1969: 68, pl. 540 (fig. 2116)) (see Fig. 20) are coarser ribbed, the ribs less
crowded and the constrictions flexuous, rather than straight. On the mature body
chamber the ribs are strong, distant, branching twice, with deep interspaces and
strong umbilical bullae. According to Collignon (1954: 30), however, the

205

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘puevynynz “py Arye

9)

‘| x ‘[ uviueduirey
O] WOT “8Z7EESD HNWA S(L68] ‘WeuUIssos]) Sisuapvdvspy (sajipvdvivy) spsaovUssoy “A-D

"SQ‘Q x ‘] uvluedues)

JO [[J UeluOJURS “puPRN[NZ “Coy AVpeo0] WOlJ “QCOIMM SVS -996I ‘uoUsI]]JOD snuussiupj)d (sajippdvavy ) SDAIIIDUSSOY “G-V ‘6l “SI

V

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

juveniles of K. (K.) karapadensis and K. (K.) besairiei grade into each other.
Kossmaticeras (Karapadites) rabenjanaharyi Collignon (1954: 33, pl. 7 (fig. 2);
1969: 69, pl. 541 (fig. 2119)) is a distinctive late form with coarser, distant ribs.

Kossmaticeras (Karapadites) hourcqui Collignon (1954: 34, pl. 10 (figs 1-2);
1969: 68, pl. 540 (fig. 2117)) has bullate umbilical nodes that give rise to strong
ribs. Kossmaticeras (Karapadites) lateconstrictus Collignon (1969: 69, pl. 541
(fig. 2122)) is in contrast characterized by dense and crowded ribs, more
numerous umbilical bullae, and striking broad constrictions on the body chamber.

Several of these species co-occur in Madagascar, and it is debatable whether
all merit specific separation. With the present poor material it is not possible to
speculate further.

The Santonian Kossmaticeras (Karapadites) planissimus Collignon, 1966
(p. 88, pl. 491 (fig. 1976)) is highly distinctive (see below), with a much more
marked loss of flank ribs on the nuclei than most later forms and a much stronger,
- coarser and more irregular ornament which readily separates it from the present
species.

Occurrence

The types are from southern India; in Madagascar the species characterizes a
Lower Campanian horizon some way above the base of the stage. The single
Zululand specimen comes from Campanian I.

Kossmaticeras (Karapadites) cf. madrasinus (Stoliczka, 1865)
Figs 23D-E

Compare

Ammonites madrasinus Stoliczka, 1865: 139, pl. 70 (figs 1-3).

Karapadites madrasinus Stoliczka: Collignon, 1954: 31, pl. 6 (fig. 5), pl. 7
(fig. 1); 1955: 305 pl. oO (fig: 5), pl) 7 (ig: 1)> pls (igs 1); 1969: 69s pleat
(fig. 2120).

Material

SAS KK105C/1, from locality 105, cliff sections 3,5 km north of the Nyalazi
River estuary, ESE of Hluhluwe, Zululand, St. Lucia Formation, imprecisely
localized in the range Santonian I1I—Campanian I.

Description

The Zululand specimen is a rather poorly preserved, composite internal
mould of the body chamber of an individual with an estimated original diameter
of circa 75 mm. The coiling appears to have been moderately involute with a
small umbilicus (estimated at approximately 30 per cent of the diameter). The
whorl section is compressed (whorl breadth to height ratio is 0,68 to 0,70), with
the greatest breadth low on the flanks, the sides slightly rounded, converging to a
narrow rounded venter.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 207

Ornament consists of numerous small comma-shaped umbilical bullae that
give rise to single ribs and pairs of ribs. These are dense, crowded, flexuous,
prorsiradiate, branch into pairs of secondaries at or about mid-flank, and are
accompanied by shorter intercalatories. There are periodic poorly preserved
constrictions.

The sutures are not decipherable.

Discussion
Of described species, this fragment compares best with Kossmaticeras

(Karapadites) madrasinus by virtue of compression of whorls, crowded ribs and
bullae. It especially resembles the specimen illustrated by Collignon (1954, pl. 7

(fig. 2)).

Occurrence

Imprecisely localized in the range Santonian IIJ—Campanian I of Zululand.
The types are from southern India. At Menabe, Madagascar, it characterizes the
Lower Campanian Karapadites karapadensis Zone, Hourcquiella bererensis
subzone.

Kossmaticeras (Karapadites) besairiei Collignon, 1954
igs 20522

Karapadites besairiei Collignon, 1954: 29, pl. 7 (fig. 3), pl. 8 (fig. 2); 1955: 28, pl. 7 (fig. 3), pl. 8
(ips 2)-1969: 68; pl. 590 (fig. 21116).

Material

SAS Z1151, from locality 105, cliff section 3,5 km north of the Nyalazi River
estuary, ESE of Hluhluwe, Zululand, St. Lucia Formation, imprecisely localized
in the range Santonian I]J—Campanian I.

Dimensions
D Wb Wh Wb:Wh U
SAS Z1151 E296, 0G00)23:5@956) 34536557) 0,83 32,063; 3)
IC 25).0 29) 34. 3(G5e7)) 0,82
at Ge (Seo (lOO) 2 38(Gies) 28-5 (C7e4) 0,84 25,3(33,4)
IC PNCO(2855) 2853674) 0,76
Description

The specimen is a largely septate internal mould retaining only a quarter of a
whorl of body chamber and traces of the original aragonitic shell.

The coiling is moderately evolute, just over 40 per cent of the previous whorl
being covered. The umbilicus comprises 33 per cent of the diameter with a
flattened, outward-sloping umbilical wall and abruptly rounded shoulder. The
whorl section is compressed (whorl breadth to height ratio varies from 0,76 to

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 20. Kossmaticeras (Karapadites) besairiei Collignon, 1954; paratype, the original of
Collignon (1954, pl. 7 (fig. 3)), from Berere, Madagascar. X 1.

0,84), with the greatest breadth at the umbilical bullae or at the shoulder in
intercostal section. The flanks are flattened and subparallel in intercostal section,
with a flattened, evenly rounded venter.

There are 20 strong comma-shaped umbilical bullae per whorl. These give
rise to pairs of, or single strong, distant rounded prorsiradiate ribs. These are
straight to feebly convex on the inner flank, feebly concave across the outer flanks
and shoulder and swing forwards across the venter, strengthening as they do so,
only to weaken over the siphonal area. Many of these ribs branch and loop across
the venter from various points on the flank while there are also shorter
intercalatories, giving a total of 55 to 60 ribs per whorl.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 209

Fig. 21. A-—C. Kossmaticeras (Kossmaticeras) aff. sakondryense Collignon, 1954; SAS Z934, from
locality 93, Zululand, Coniacian II. D-E. Kossmaticeras (Karapadites) planissimus Collignon,
1966; SAS KK105B, from locality 105, Zululand, Santonian III or Campanian I. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

210

IO JJ] ueUOJUeS ‘puelniNZ ‘coy

Ayyesoy wory

‘I X ‘[ uewedues)
‘ISLIZ SVS ‘pS6l ‘uousI]JOD 1amiwsaq (sanpodvavy) spsavdyvUssoy

‘7 ‘SI

CRETACEOUS FAUNAS FROM SOUTH AFRICA Di

There are six strong, broad and deep constrictions per whorl on the mould,
flanked by collar-ribs. The adapertural collar is usually simple, with a weak or no
umbilical bulla. The adapical collar splits into three secondary virgatotome ribs.

The suture-line is as in Kossmaticeras (Karapadites) planissimus, described
below.

Discussion

This magnificent specimen compares well with the holotype (Collignon 1954,
pl. 8 (fig. 2)), and the body chamber paratype (Collignon 1954, pl. 7 (fig. 3)),
reillustrated here as Figure 20, showing the same distinctive strong bullae, distant
ribs, effaced on the inner flank but strong on outer flank and venter, and similar
constrictions and associated collar-ribs. The variety Kossmaticeras (Karapadites)
planissimus bererensis Collignon (1954: 31, pl. 9 (fig. 1)) is even more coarsely
and strongly ribbed.

Occurrence
Lower Campanian Kossmaticeras (Karapadites) besairiei Zone of Menabe,

Madagascar, especially the Hourcquiella bererensis subzone. Santonian III or
Campanian I of Zululand.

Kossmaticeras (Karapadites) planissimus Collignon, 1966

Figs 1A, 19A—B, 21D-E, 23A—-C, 24B, 25-26
Karapadites planissimus Collignon, 1966: 38, pl. 541 (fig. 1976).

Types

The holotype, by original designation, is the original of Collignon (1966: 88,
pl. 541 (fig. 1976)), from the Upper Santonian Pseudoschloenbachia umbulazi
Zone of Collignon’s (1969) locality 692, Ampamba-Antsirasira (Belo-sur-
Tsiribihina), Madagascar. There are 11 other, unfigured paratypes.

Material

SAS KK105, 105B, Z1954 and SAS H126 A/3, from locality 105, cliff section
3,5 km north of the Nyalazi River estuary, St. Lucia Formation, Santonian III or
Campanian I, Zululand.

YPM 1071, from ‘Port Natal’—the specimen shows signs of water wear, and
may be from the Umzamba Formation of southern Natal or of the Transkei.

Dimensions

D Wb Wh Wb:Wh wi
SAS Z1954 104,0(100) 36,5(35,1) 40,8(39,2) 0,90 25,8(34,4)
SAS KK105B 113,7(100) 38,0(33,4) 43,5(38,2) 0,87 36,4(32,0)
SAS H126A/3 114,0(100) 36,0(31,5) 43,2(37,9) 0,83 36,5(32,0)

SAS KK105 141,0(100) 42,5(30,1) 51,0(36,2) 0,83 46,5(33,0)

212 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. A-—B. Kossmaticeras (Karapadites) planissimus Collignon, 1966; SAS Z1954.
C-E. Kossmaticeras (Karapadites) cf. madrasinus (Stoliczka, 1865); SAS KK H105C/1. Both
from locality 105, Zululand, Santonian IIT or Campanian I. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 213

UD oL

—

Fig. 24. External sutures of: A. Kossmaticeras (Karapadites) karapadensis
(Kossmat, 1897), BMNH C83328; B. Kossmaticeras (Karapadites) planissimus
Collignon, 1966, SAS KK105B. Both x 2.

Description

The material available is generally well preserved and in the form of both
moulds and specimens retaining the original aragonitic shell.

SAS Z1954 shows the early growth stages, at a diameter of 18,5 mm
(Fig. 23A-B). The coiling is moderately involute (48 per cent of the previous
whorl is covered), the whorl section depressed (whorl breadth to height ratio is
1,1). Ornament consists of umbilical bullae only on the internal mould (the shell
is not preserved at this diameter). There are four to five strong constrictions per
half whorl. They are broad, deep, prorsiradiate, straight on the flanks, and pro-
jected forward over the ventro-lateral shoulders into a narrowly rounded ventral
convexity. Each has a narrow rib on the adapertural side, and this is followed by a
second, shallower constriction. A highly distinctive growth-stage follows this and
extends to the beginning of the outer whorl at an estimated diameter of 60 mm.
The umbilicus is thus shallow, the umbilical wall slopes outward and is flattened,
with an abruptly rounded umbilical shoulder. All those parts of the flanks not

ANNALS OF THE SOUTH AFRICAN MUSEUM

214

‘AUSIOAIU JVA ‘wWinasnyy Apoqeag oy} ut

"CL'0 X (| [PIWN Od, Woy
ILOL ON Uoumtdeds ‘qq6] ‘uoUsTJOD snuussiunjd (sajpodvavy) sviaoyvwussoy “SZ “B14

CRETACEOUS FAUNAS FROM SOUTH AFRICA 215

Fig. 26. Kossmaticeras (Karapadites) planissimus Collignon, 1966; specimen No 1071 in the
Peabody Museum, Yale University, from ‘Port Natal’. x 1.

concealed by the succeeding whorls are flat. There are 16 to 19 strong comma-
shaped umbilical bullae of variable strength that give rise to single or paired,
markedly prorsiradiate ribs. These decline markedly on the inner to mid-flank
which in some specimens (e.g. YPM 1071, see Figs 25, 26) are almost smooth.
The ribs flex back across the outer flank, where they are concave and

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

accompanied by intercalated ribs and may branch before sweeping forward across
the venter, where they are at their strongest. There are up to eight constrictions
per whorl.

All the available specimens are adult, with diameters of up to 140 mm and up
to two-thirds of a whorl of body chamber. At mature growth-stages, at shell
diameters exceeding 60 mm, the coiling becomes increasingly evolute and the
umbilicus widens to comprise up to 34 per cent of the diameter. The umbilical
wall is of moderate height, is flattened, and slopes outward, giving a shallow
crater-like form to the adult umbilicus. The whorls are compressed (whorl
breadth to height ratio is as little as 0,83), with the greatest breadth at the
umbilical bullae. The flanks are compressed, subparallel to slightly convergent
with a broadly rounded venter. There are up to 20 strong to weak, variable
umbilical bullae per whorl. These give rise to single, or pairs of, ribs on the last
part of the phragmocone and predominantly single ribs on the body chamber. The
~ ribs are strong, broad, rounded, prorsiradiate and gently flexuous, pass straight
across the inner flank, are convex at mid-flank, and sweep back across the outer
flank, where they are concave, before sweeping forward to pass across the venter
with a slight convexity. Some ribs strengthen markedly and branch high on the
flank, looping across the venter, while there are occasional intercalated ribs,
giving a total of 60 per whorl.

SAS H126A/3, KK105 and YPM 1071 all show the adult aperture, which is
preceded by a final section of shell ornamented by dense, simple, fine flexuous
ribs and growth striae, lacking bullae. The mouth border itself appears to have
been simple.

Adult growth-stages appear to have had five or six constrictions per whorl,
but these are much less conspicuous and relatively shallower than on the inner
whorls, especially where the shell is lacking. The associated collar-ribs are,
however, highly distinctive. The adapical one is strong, bullate, and bifurcates
over the venter, and the adapertural branch develops a much more narrowly
rounded convexity than the adapical branch. The adapertural collar is, by
contrast, much weaker, simple, unbranched, narrower, and lacks a bulla.

The suture-line is shown in Figure 24B.

Discussion

Kossmaticeras (Karapadites) planissimus is the only species of the subgenus
so far recorded from the Santonian. Flat sides, feeble flank ornament when young
plus very coarse, distant ribs of irregular length and branching make it
immediately distinctive and easily separated from all other species.

Occurrence

Upper Santonian, Pseudoschloenbachia umbulazi Zone of Menabe,
Madagascar; Santonian III or Campanian I of Zululand and probably also the
Transkei (Umzamba Formation) where it is of Middle Santonian or Lower
Campanian age.

CRETACEOUS FAUNAS FROM SOUTH AFRICA DAG.

Genus Maorites Marshall, 1926

Type species
Kossmaticeras tenuicostatum Marshall, 1917: 445, text-fig. 3, pl. 33 (fig. 1).

Maorites cf. subtilistriatus Collignon, 1954
Figs 27—28

Compare

Maorites subtilistriatus Collignon, 1954: 38, pl. 11 (fig. 3); 1969: 72, pl. 442
(fig. 2124).

Types

The holotype is the original of Collignon (1954, pl. 11 (fig. 3)), from the
Lower Campanian of Berere, Madagascar, refigured here as Figure 29. There are
two unfigured paratypes (Collignon 1954: 38).

Material

Two specimens only: SAS Z709 from the Nibela Peninsula, Zululand,
St. Lucia Formation, probably Campanian II or II]; BMNH C83332 from bed 18,
locality 110 on the Nibela Peninsula, St. Lucia Formation, Campanian III.

Dimensions

D Wb Wh Wb:Wh U
SAS Z709 at 134,0 43,2(32,2) 62,3(46,5) 0,69 S09 Ose)
Description

The two specimens are rather poorly preserved. BMNH C83332 is largely
septate with an estimated maximum preserved diameter of 105 mm. It is in part
an internal mould, in part bearing recrystallized and overgrown shell that hides
much of the detail of the ornament. SAS Z709 is similarly overgrown but appears
to be adult, with more than half a whorl of body chamber. The estimated adult
diameter must have approached 160 mm.

The coiling is moderately involute, about 60 per cent of the previous whorl
being covered. The umbilicus comprises 23 per cent of the diameter and is of
moderate depth; the umbilical wall is flattened and at 90 degrees to the flank of
the preceding whorl. The whorls are compressed (whorl breadth to height ratio is
0,69). The greatest breadth is at the umbilical shoulder, from which the flattened
flanks converge slightly to a broadly rounded, somewhat flattened venter.

On BMNH C83332, the internal mould shows an ornament of dense,
flexuous prorsiradiate ribs. On the test, where preserved, they are stronger, with
steep sides and distinctly flattened tops. They are narrow at the umbilical
shoulder but broader and flatter across the flanks and ventrolateral shoulders. On
the venter, which they pass straight across, they are at their strongest

ANNALS OF THE SOUTH AFRICAN MUSEUM

218

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CRETACEOUS

FAUNAS

FROM SOUTH AFRICA

219

III. x 1.

lan

; BMNH C83332, from locality 110, Zululand, Campan

Fig. 28. Maorites cf. subtilistriatus Collignon, 1954

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

development; they are weaker and rounded on the mould but flattened and bar-
like, with narrower slot-like interspaces, where the test is preserved. They
increase progressively in strength as the diameter increases in the smaller
specimen but on the body chamber of SAS Z709 they show a marked
strengthening over the last quarter whorl. There is occasional branching at
various points on the flank.

There are periodic narrow, flexuous, prorsiradiate constrictions that are most
obvious on the internal mould, where they are strongest over the umbilical
shoulder. In BMNH C83332 there are an estimated ten per whorl, associated with
broad collars on the mould. It is not clear whether or not they truncate ribs. In
SAS Z709 constrictions are again well developed and appear to be associated with
stronger collars where the shell is preserved.

Fig. 29. Maorites subtilistriatus Collignon, 1954; the holotype, the original
of Collignon (1954, pl. 9 (fig. 3-3a)), from the Lower Campanian of Berere,
Madagascar. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 221

The suture-line is partially exposed on BMNH C83332, and is deeply and
intricately subdivided.

Discussion

Howarth (1966) has suggested that Maorites subtilistriatus from Madagascar
(and now Zululand), and M. tenuicostatus from New Zealand (see Henderson
1970: 50, pl. 9 (figs 3-4), pl. 10 (fig. 2)) are synonyms of the equally finely-ribbed
M. densicostatus (Kilian & Reboul, 1909) (p. 30, pl. 15 (fig. 4), pl. 18 (fig. 1))
from Antarctica. Henderson (1970) points out that M. tenuicostatus retains fine
ribs to a large size, but admits that juvenile M. tenuicostatus and M. densicostatus
are probably inseparable. Our material is too poor to resolve the problem, so we
use the Madagascan name, although suspecting that but a single full species is
present. Indeed, so subtle are intraspecific differences in Maorites that, given
large populations and clear recognition of dimorphism, there can be little doubt
that only a few species could be reasonably maintained.

Maorites multiconstrictus Henderson, 1970 (p. 51, pl. 9 (fig. 2)) is
distinguished by coarse flexuous ribs with far more constrictions, and distinctive
ontogenetic changes. Maorites angulocostatus Henderson, 1970 (p. 52, pl. 10
(fig. 1)) has much stronger bullae and distant, sickle-shaped fine ribs. Maorites
mackayi (Hector, 1886) (Henderson 1970: 53, pl. 10 (fig. 3)) is a poorly known
species most easily recognized by the wide spacing of the fine ribs. Maorites
seymourianus (Kilian & Reboul, 1909) (p. 29, pl. 19 (fig. 1)) is distinguished most
readily by the stronger umbilical bullae. Maorites menabensis Collignon, 1954
(p. 37, pl. 11 (fig. 2)) is a rather broad-whorled species with regularly dichoto-
mous ribs. Maorites tuberculatus, Howarth, 1958 (p. 11, pl. 2 (figs 1-3)) 1s
immediately distinguishable by the strongly rounded whorls, deep, broad
constrictions and large umbilical bullae. Maorites pseudobhavani Spath, 1953
(p. 25, pl. 6 (figs 7-9)) is an evolute, robust but diminutive species with strong
umbilical bullae that give rise to groups of ribs; it should be referred to Gunnarites
(fide Howarth 1966: 67). Maorites kandi (Stoliczka, 1865) (p. 140, pl. 70 (fig. 4))
is more evolute with lower, slowly expanding whorls. The ribs are coarser, and
show marked irregularity and become widely spaced at the aperture. Maorites
aemilianus (Stoliczka, 1865) (p. 141, pl. 70 (figs 6—8)) is characterized by elongate
bullae, and the ornament of fine ribs arranged in bundles is distinctive. Maorites
magnumbilicatus Collignon, 1954 (p. 40, pl. 12 (fig. 1)) is a more evolute,
massively whorled species with a deep umbilicus; ornament is initially of fine,
crowded, slightly flexuous ribs which become strong and straight on the beginning
of the body chamber, thereafter disappearing to leave the greater part of the body
chamber smooth, according to Collignon (1954: 40).

Occurrence

Maorites subtilistriatus characterizes the Maorites aemilianus subzone of the
Karapadites karapadensis Zone in the Lower Campanian of Menabe, Madagascar.

VD? ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Gunnarites Kilian & Reboul, 1909

Type species
Olcostephanus antarcticus Weller, 1903: 4, by the subsequent designation of
Diener (1925: 101).

Gunnarites antarcticus (Weller, 1903)

Figs 30-33, 34D-E

Olcostephanus antarcticus Weller, 1903: 4, pl. 2 (figs 1-2).

Gunnarites antarcticus Stephen Weller: Diener, 1925: 101 (with synonymy). Spath, 1953: 29, pl. 3
(fig. 5), pl. 4 (fig. 9), pl. 6 (figs 1-2, 4-5), pl. 11 (fig. 1). Wright 1957: 374, fig. 490 (4).
Howarth, 1966: 66 et seq.. Lahsen & Charrier, 1972: 529, pl. 1 (figs 4-6).

Gunnarites antarcticus Weller, var. monilis Spath, 1953: 31, pl. 6 (fig. 3).

Gunnarites antarcticus Weller, var. inflata Kilian & Reboul: Spath, 1953, pl. 7 (fig. 1), pl. 8
(fig. 8).

Gunnarites gunnari Kilian & Reboul: Spath, 1953: 33, pl. 5 (figs 4-5) (with synonymy).

Gunnarites pachys Spath, 1953: 34, pl. 9 (figs 1-3) (including var. media).

Gunnarites flexuosus Spath, 1953: 35, pl. 3 (figs 3-4), pl. 9 (figs 4-5).

Gunnarites rotundus Spath, 1953: 36, pl. 12 (figs 1-3) (including varieties kalikaformis and
compressa).

Gunnarites paucinodatus Spath, 1953: 37, pl. 7 (fig. 4).

Gunnarites aff. G. antarcticus (St. W.): Blasco de Nullo, Nullo & Proserpio, 1980: 487, pl. 5
(figs 9-10).

Material

BMNH C83336, from the St. Lucia Formation, Campanian III at locality
115; BMNH C83334, St. Lucia Formation, Maastrichtian a (= ‘Campanian’ IV),
locality 113. SAS Z224/1 and an unregistered and unlocalized specimen in the
South African Geological Survey Collections are also referred to the species.

Dimensions

D Wb Wh Wb: Wh U
SAS 2224/1 130,0 38,8(29,8) 55,0(42,3) OI 40,3(31)
BMNH C83334 108,5 36,3(33,5) 45,5(41,9) 0,80 36;5(3356)
Description

The best-preserved specimen is BMNH C83334, represented by a well-
preserved external mould, a whorl of septate phragmocone and the beginning of
the body chamber. Coiling is moderately involute, 56 per cent of the previous
whorl being covered. The umbilicus comprises between 31 and 33,6 per cent of
the diameter and is of moderate depth with a subvertical wall. The whorl section
of all our specimens is compressed (breadth to height ratio varies from 0,71 to
0,80, with the greatest breadth at, or close to the umbilical bulla). The whorl sides
are flattened and convergent, the venter broadly and evenly rounded.

There are 18 strong, sharp umbilical bullae per whorl, projected into the
umbilicus. These give rise to groups, generally of three ribs, while one or two

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CRETACEOUS FAUNAS FROM SOUTH AFRICA

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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CRETACEOUS FAUNAS FROM SOUTH AFRICA Tes

Fig. 32. Gunnarites antarcticus (Weller, 1903); BMNH C83334, from locality 115,
Zululand, Maastrichtian a. X 1.

non-bullate ribs extend to the umbilical shoulder between these groups. Shorter,
intercalated ribs arise around mid-flank. The ribs are initially narrow, but
broaden over the venter, are flat-topped, prorsiradiate and straight or feebly
flexuous, totalling over 60 per whorl. All are strongly denticulate, the spiral
denticulations most prominent on the shell over the ventrolateral and ventral
regions.

There are six narrow, deep constrictions per whorl, preceded by a thickened
rib, and followed by a broad interspace, sometimes with associated rather feeble
riblets.

The deeply incised suture-line (Fig. 31) is typically kossmaticeratid.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Gunnarites antarcticus (Weller, 1903); BMNH C83336,
from locality 115, Zululand, Maastrichtian a. x 1.

Discussion

Howarth (1966) has pointed out the intergrading relationship between all the
Antarctic species of Gunnarites described by Spath (1953), and examination of
the Antarctic material suggests there are only two forms, the large G. antarcticus
and small G. kalika (Stoliczka, 1865). We strongly suspect these to be dimorphs,
but cannot fully prove it at this time, so that they are maintained as separate here.

Of the various forms described from Antarctica, the specimens described here
most closely recall the specimen figured by Spath (1953) as his plate 4 (fig. 9a—b).

Gunnarites antarcticus differs very clearly from the various New Zealand
species described and discussed by Henderson (1970). Thus G. zelandicus

Fig. 34 (facing page). .A—C. Gunnarites kalika (Stoliczka, 1865). A.SAM-—PCO5907,

from the offshore Alphard Group (figured by Klinger, Kauffman & Kennedy, 1980,

fig. 6A-B). B—C. An unregistered specimen in the South African Geological Survey

Collections. D—E. Gunnarites antarcticus (Weller, 1903), an unregistered specimen in the
South African Geological Survey Collections. All x 1.

CRETACEOUS FAUNAS

FROM SOUTH AFRICA

fei

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Marshall, 1917) (see Henderson 1970: 54, pl. 11 (fig. 1)) has only eight umbilical
bullae and four constrictions per whorl. Gunnarites denticulatus (Marshall, 1917)
(p. 55, pl. 11 (fig. 1), pl. 12 (fig. 2)) is much more densely and delicately ribbed.
Gunnarites spathi Henderson, 1970 (p. 56, pl. 12 (figs 1—-4)) is a sparsely ribbed
species in the same relationship to G. antarcticus as Kossmaticeras sparsicostatum
is to K. theobaldianum. Gunnarites varicostatus Henderson, 1970 (p. 57, pl. 13
(figs 1, 3)) is an evolute form with markedly prorsiradiate and flexuous ribs.

Occurrence

Campanian III and Maastrichtian a (‘Campanian’ IV) and Maastrichtian b of
Zululand. The Antarctic occurrences are variably dated as Upper Campanian
(Spath 1953), Lower—Middle Campanian (Howarth 1966) or uppermost Campa-
nian—basal Maastrichtian (Henderson 1970). Also the Maastrichtian of
_Magellanes Province, Chile.

Gunnarites kalika (Stoliczka, 1865)
Fig. 34A—C

Ammonites kalika Stoliczka, 1865: 140, pl. 70 (fig. 5).

Holcodiscus kalika (Stoliczka): Kossmat, 1898: 41 (148).

Gunnarites kalika (Stoliczka): Kilian & Reboul, 1909: 34; Spath, 1953: 33, pl. 10 (figs 1-6);
Howarth, 1966: 16; Blasco de Nullo, Nullo & Proserpio, 1980: 487, pl. 4 (figs 2-4); Lahsen
& Charrier, 1972: 529, pl. 2 (figs 3-6).

Maorites pseudobhavani Spath, 1953: 25, pl. 6 (figs 7, 9 non 8?); non pl. 11 (figs 2-4).

Gunnarites cf. kalika (Stoliczka: 1865): Klinger, Kauffman & Kennedy, 1980: 299, fig. 6A—-B.

Material

A single unregistered and unlocalized specimen in the South African
Geological Survey Collections, from the Campanian—Maastrichtian St. Lucia
Formation on the western shores of Lake St. Lucia. SAM—PCO5907, offshore
Alphard Group, South Africa.

Dimensions

D Wb Wh Wb: Wh U
SAS Dyes) IeS(GOR0) 2580 (43ers) 0,69 17,4(60,2)
Description

The specimen is a composite mould and is adult, with approximately two-
thirds of a whorl of body chamber. Coiling appears to have been moderately
involute, the umbilicus comprising 30 per cent of the diameter. The whorl section
is compressed (whorl breadth to height ratio is 0,69) with the greatest breadth low
on the flank or at the umbilical bulla.

The flanks are high, flattened, and converge to a narrow, rounded, flattened
venter. Sharp umbilical bullae project into the umbilicus and give rise to bundles
of up to four primary ribs. Between bullae pairs of ribs or single ribs arise at the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 229

umbilical shoulder; all ribs are crowded, narrow, rounded, prorsiradiate and
gently flexuous, occasionally branching low or high on the flanks, without
conspicuous intercalatories. The ribs thicken over the venter (which then pass
straight across) and are distinctly denticulate.

There are five constrictions per half whorl. These are relatively deep,
prorsiradiate and feebly flexuous. They are flanked by collar-ribs; the adapical
one is the stronger, arising at an umbilical bulla and branching twice; once low on
the flank and again at the ventrolateral shoulder.

There is a distinctive change in ornament over the last part of the body
chamber with the ribs becoming finer and much more flexuous.

Discussion

This beautiful specimen appears to be adult, as is shown by the loosening of
coiling and feeble ornament developed at the end of the body chamber. It
matches well with both the holotype and other specimens cited in the synonymy.

In both South Africa and Antarctica, this small form occurs with the much
larger Gunnarites antarcticus, discussed above, and we strongly suspect that it
may be the microconch of that species, but cannot prove it. The two species are
thus left separate at present.

Gunnarities kalika differs from G. zelandicus (Marshall, 1917) (see Hender-
son 1970: 54, pl. 11 (fig. 1)), G. spathi Henderson (1970: 56, pl. 12 (figs 1, 4)) and
G. varicostatus Henderson (1970: 57, pl. 13 (figs 1, 3)) in being compressed,
involute, high-whorled and delicately ribbed, and from G. denticulatus (Marshall,
1926) (Henderson 1970: 55, pl. 11 (fig. 2), pl. 12 (fig. 2)) in its more prominent
constrictions and lack of prominent bullae.

Occurrence

This species is known from southern India, Antarctica, Patagonia, Zululand
and the offshore Alphard Group. The offshore specimen occurs with Eubaculites
latecarinatus (Brunnschweiler, 1966), suggesting an early Maastrichtian age—
Maastrichtian I in the sense of Kennedy & Klinger (1975). The Antarctic
specimens are regarded as largely Upper Campanian (Spath 1953), Lower to
Middle Campanian (Howarth 1966: 68) or uppermost Campanian—basal Maas-
trichtian (Henderson 1970: 78). The holotype is from Ootacod, southern India, a
locality yielding undoubted Maastrichtian species. In Patagonia it is said to be
Upper Maastrichtian.

ACKNOWLEDGEMENTS

We thank Dr C. W. Wright (Oxford), Dr M. K. Howarth and Mr D. Phillips
(British Museum (Natural History), London), Drs C. K. Brain and E. Vrba
(Transvaal Museum, Pretoria), and the staff of the Geological Collections,
University Museum, Oxford and South African Museum, Cape Town for their

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

advice and assistance. The financial support of the Sir Henry Strakosch Bequest
and the Natural Environment Research Council to Kennedy is gratefully
acknowledged, as is support from the Council for Scientific and Industrial
Research to Klinger.

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6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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WILLIAM JAMES KENNEDY
&
HERBERT CHRISTIAN KLINGER

CRETACEOUS FAUNAS FROM
ZULULAND AND NATAL, SOUTH AFRICA
THE AMMONITE FAMILY
KOSSMATICERATIDAE SPATH, 1922

95 PART 6 ~— JUNE 1985 ISSN 0303-2515

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

Fiscuer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, ese masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4 ): $1.

THIELE, J. 1910. Mollusca: B. Polypiseopliont Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270

(continued inside back cover)

ENNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
June 1985 Junie
Part 6 Deel

THE AEOLID NUDIBRANCH
FAMILY AEOLIDIITDAE
(GASTROPODA, OPISTHOBRANCHIA)
FROM TROPICAL SOUTHERN AFRICA
By
TERRENCE M. GOSLINER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

Fig. 1. Living animals. A. Aeolidiella alba Risbec. 1928.
B. Baeolidia palythoae sp. nov. C. Berghia chaka sp. nov.

THE AEOLID NUDIBRANCH FAMILY AEOLIDITDAE
(GASTROPODA, OPISTHOBRANCHIA)
FROM TROPICAL SOUTHERN AFRICA

By
TERRENCE M. GOSLINER

National Museum of Natural History,
Smithsonian Institution, Washington, D.C.*

(With 18 figures and 4 tables)

[MS accepted 27 June 1984]

ABSTRACT

Three species of nudibranchs belonging to the Aeolidiidae are recorded from the shores of
southern Africa for the first time. Aeolidiella alba has previously been recorded from other
localities in the Indo-West Pacific and Panamic regions. Baeolidia palythoae sp. nov. and Berghia
chaka sp. nov. are described. Difficulties in producing a natural classification of the Aeolidiidae
are discussed and the status of several species within the genera Aeolidiella, Aeolidiopsis,
Baeolidia, Berghia, and Spurilla is altered.

CONTENTS

PAGE

COE RO/CMY CVO) Os 3 ee Mee lean eee cee a LAR, SARE ZO ee 233
SSSR ITIUICINS te ee Meee ee ae Sans ME eRe ea. Dee Re 234
CONGUE OMA ISDEC) VOI «putea. =p aeheee es Be ee 234
BACOMAA DAV ULOGE SD: NOMS =) so et a eee Di,

iCVOP OVO G IVOUITIES Oy 10) ree see ER ar Regn ie 245
DISCUSSIONS RS Bh a Ste oe ae ae ene Semen ED Ea 8 at 250
(SCMCMCGSIOMIVASIOM ns ol Sak ne a oe hae De Phe oe ee 250

Key iothe genera of the Acolidudae .) 4.223802. -7 22: 759
Discussion of Baeolidia palythoae sp.nov. ............. 262
DISCUSSION Ol Berenia Chaka sp: DOV..2 52255222. 5-25-5- 262
POON EGSCIMEMIS: 46 as sno. oe auiie ou ae Be Hise ena ere ae 265
Ree INC ity ee a te Ica Ea ee cil Bw eu Racin aul ORNS 265

INTRODUCTION

The aeolidacean nudibranchs of southern Africa have only been superficially
studied (Bergh 1907; Thiele 1925; Barnard 1927). More recent studies (Macnae
1954; Gosliner & Griffiths 1981) have focused specifically upon the aeolidacean
fauna and have recorded several additional taxa from the region. All of these
studies have dealt primarily with the temperate regions of the Cape Province of
South Africa. There are no records of any aeolidacean nudibranchs from the
subtropical and tropical portions of the region.

* Present address: Department of Invertebrate Zoology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA.

239

Ann. S. Afr. Mus. 95 (6), 1985: 233-267, 18 figs, 4 tables.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Recent collection of opisthobranch gastropods from Transkei and Natal
waters has yielded specimens of many taxa that have not previously been
recorded from southern Africa, including three species of the family Aeolidiidae.
It is the intent of this paper to describe the morphology of these species and to
discuss their systematic placement.

Specimens have been deposited in the following institutions: South African
Museum, Cape Town (SAM); United States National Museum of Natural
History, Washington, D.C. (USNM); and California Academy of Sciences
(CASIZ).

DESCRIPTIONS
Aeolidiella alba Risbec, 1928

Bigs 1Ay 2,3

‘Aeolidiella alba Risbec, 1928: 261, fig. 87, pl. 10 (fig. 9).
Spurilla alba (Risbec, 1928) Edmunds, 1969: 465, fig. 9.

Material

SAM-—A35648: 1 specimen; Salt Rock, Natal; intertidal zone; 30 April 1981;
collected by T. M. Gosliner. SAM—A35643: | specimen; Jesser Point, Sodwana
Bay National Park, Natal; intertidal zone; 19 May 1981; collected by T. M.
Gosliner. CASIZ 055326: 5 specimens; Jesser Point, Sodwana Bay National Park,
Natal; intertidal zone; 6 May 1982; collected by T. M. Gosliner. Uncatalogued:
1 specimen; Adlam’s Reef, Sodwana Bay National Park, Natal; intertidal zone;
7 May 1982; collected by T. M. Gosliner. 2 specimens; Jesser Point, Sodwana Bay
National Park, Natal; intertidal zone; 7 May 1982; collected by T. M. Gosliner.

Distribution

Widespread in the Indo-West Pacific: Tanzania (Edmunds 1969); Australia
(Burn 1966); New Caledonia (Risbec 1928); Nayarit and Sonora, Mexico (Sphon
1971, 1978). Recently also reported from the Atlantic (Barbados) by Edmunds &
Just (1983).

External morphology

The preserved animals range from 4 to 7 mm in length. The oral tentacles are
short and cylindrical, often with a swelling near their middle. The rhinophores are
longer than the oral tentacles and possess a pair of bulbous swellings in their outer
half. The foot (Fig. 2A) is expanded near its anterior limit but there are no
distinct foot corners. The foot is broad throughout its length. The somewhat
dorso-ventrally flattened cerata are arranged in diagonal linear rows. There are
four ceratal rows in the anterior right digestive branch with five or six rows in the
posterior right branch. The ceratal rows of the anterior branch each contain from
seven to ten cerata per row, while the posterior rows contain one to seven cerata.
The gonopore is situated near the antero-ventral base of the second ceratal row.
The anus is located immediately posterior to the fifth ceratal row.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 235

Fig. 2. Aeolidiella alba Risbec, 1928. A. Ventral view. Scale = 1,0 mm.
B. Buccal region. Scale=1,0 mm. C. Jaw. Scale =0,5 mm.
D. Reproductive system. Scale = 0,5 mm.

Coloration

The living animals (Fig. 1A) are translucent white covered with opaque white
on the notum, head, oral tentacles and cerata. The rhinophores and their bases
are translucent orange. At the base of each ceras is a dark brown glandular area.

Digestive system

A pair of large oral glands (Fig. 2B), consisting of large vesicles, extends
posteriorly from their openings near the mouth to well beyond the buccal mass.
The jaws (Fig. 2C) are strong with an elongate masticatory border that is devoid

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

ASR Bo]

Fig. 3. Aeolidiella alba Risbec, 1928. Scanning electron micrographs of radula.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 237

of denticles. The uniseriate radula (Fig. 3) consists of 19-27 pectinate teeth with
21-30 elongate denticles on either side of the prominent central cusp.

Reproductive system

The reproductive system (Fig. 2D) occupies most of the body cavity posterior
to the buccal mass. The ovotestis consists of numerous lobules, each containing
distinct male and female acini. The ampulla is narrow and divides into a short
oviduct and an elongate vas deferens. By means of an elongate duct the spherical
receptaculum seminis joins the ampulla at its bifurcation into male and female
ducts. The oviduct empties into the albumen gland. Adjacent to the albumen
gland is the membrane gland. The mucous gland is bilobed and forms the bulk of
the reproductive organs. It terminates at a ventral gonopore. The vas deferens
expands into a prostatic portion, terminating at the tip of the unarmed penis. The
separate male gonopore is dorsal to the female gonopore.

Natural history

Aeolidiella alba has been found in association with small white acontiate sea
anemones, on which it presumably feeds. All of the South African specimens
have been observed in small rock pools in the mid- and lower intertidal zone. The
animals are active at night, when they are often seen at or near the surface of rock
pools. During the day they have been found under small rocks or dead pieces of
coral.

Baeolidia palythoae sp. nov.
Figs 1B, 4-10

Material

Holotype. SAM-—A35640: Umgazana, Transkei; intertidal zone; 23 April
1982; collected by T. M. Gosliner.

Paratypes. SAM-—A35636: 2 specimens; Jesser Point, Sodwana Bay National
Park, Natal; intertidal zone; 9 May 1981; collected by T. M. Gosliner. SAM-—
A35638: 2 specimens; Jesser Point, Sodwana Bay National Park, Natal; intertidal
zone; 6 May 1982; collected by T. M. Gosliner. SAM—A35639: 4 specimens;
Adlam’s Reef, Sodwana Bay National Park, Natal; intertidal zone; 9 May 1981;
collected by M. Cooke. SAM-—A35641: 2 specimens; Ramsgate, Natal; intertidal
zone; 9 March 1981; collected by T. M. Gosliner. SAM—A35644: 3 specimens;
Park Rynie, Natal; intertidal zone; 10 March 1981; collected by T. M. Gosliner.
SAM-—A35646: 1 specimen; Adlam’s Reef, Sodwana Bay National Park, Natal;
9 May 1981; collected by M. Cooke. USNM 805051: 1 specimen; south side of
St. Anne Channel, Seychelles Islands; depth 3 m; 11 December 1964; collected
by L. Pierce. CASIZ 053777: 3 specimens (one partially dissected); Jesser Point,
Sodwana Bay National Park, Natal; intertidal zone; 9 May 1981; collected by
T. M. Gosliner. CASIZ 053778: 2 specimens; Jesser Point, Sodwana Bay National
Park, Natal; 6 May 1982; collected by T. M. Gosliner.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The name palythoae is derived from the genus of zoanthid anthozoans,
Palythoa, on which this species feeds.

Distribution

Specimens have been collected along the coast of southern Africa from
Umgazana, Transkei (31°43’S 29°25'E), to Jesser Point, Sodwana Bay National
Park, Natal (27°32'S 32°41’E). A single specimen in the collections of the
National Museum of Natural History was collected from the Seychelles Islands.

External morphology

Living animals (Fig. 1B) may reach a length of 17 mm. The oral tentacles are
short and slender, tapering near their apices. The rhinophores (Fig. 4A) are
approximately equal in length to the oral tentacles and are sparsely covered by
- elongate tubercles. The foot (Fig. 4B) is broadest anteriorly and simply rounded,
without angular or tentacular extensions. The anterior margin of the foot is
weakly grooved. The cerata (Fig. 4D) are dorso-ventrally flattened and broadly
ovoid in shape. There is a single ceratal row in the right anterior digestive branch
(Fig. 4C). Behind the interhepatic space are the four to six ceratal rows of the
right posterior digestive system. There are from three to five cerata per ceratal
row in the anteriormost rows. The posterior ceratal rows contain one to three
cerata per row. The gonopore is situated below the middle of the first ceratal row.
The anus is located postero-ventrally to the second ceratal row.

Coloration

The ground colour is translucent yellowish. The yellow colour is overlaid
with a reticulate brown pattern, which varies in its density. The tip of each ceras
bears a small opaque white spot. Ventrally and slightly eccentric to the apex is a
larger spot of dark brown pigment. This pattern of coloration did not vary in any
of the approximately 100 specimens of this species observed.

Digestive system

There are two pairs of oral glands present in the buccal region (Fig. 5A). The
larger pair inserts into the buccal mass on its antero-dorsal side. These glands are
elongate and may be convoluted. They extend posteriorly at least twice the length
of the buccal mass. The smaller pair of glands is situated more anteriorly. They
are ovoid in shape and each empties on the ventral side of the head (Fig. 4B) by
means of a slit-like pore, which is readily visible in all living and preserved
material.

The jaws (Figs 5B, 7, 8A) are moderately strong and narrow. The elongate
masticatory border is devoid of denticles but bears numerous small tubercles
along the inside of its entire length (Figs 7B, 8B).

The radula is uniseriate with 17-23 teeth. The teeth (Figs 8B, 9) are evenly
curved with 41-55 denticles along either half of the tooth. A central denticle is
absent.

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eS)
No)

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

oa
CN
m
O
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0
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Fig. 4. Baeolidia palythoae sp. nov. A. Rhinophore. Scale = 0,5 mm.
B. Ventral view. Scale=1,0 mm. C. Lateral view. Scale=4,0 mm. D. Ceras.
Scale = 1,0 mm.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Baeolidia palythoae sp. nov. A. Buccal region. Scale = 2,0 mm. B. Jaw.
Scale = 0,25 mm.

Fig. 6. Baeolidia palythoae sp. nov. A. Central nervous system. Scale = 1,0 mm.
B. Reproductive system. Scale = 1,0 mm.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

1SKY WO: 37MM 8 86$:66608 P: 66034

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Fig. 7. Baeolidia palythoae sp. nov. Scanning electron micrographs.
B. Masticatory border of jaw.

Aw Jaw:

VAD ANNALS OF THE SOUTH AFRICAN MUSEUM

l 4 1SKY WO: 12MM 8 86$:66006 P: 86005

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Fig. 8. Baeolidia palythoae sp. nov. Scanning electron micrographs. A. Detail of
masticatory papillae. B. Radular teeth.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

Fig. 9. Baeolidia palythoae sp. nov. Scanning electron micrographs of radular teeth.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Baeolidia palythoae sp. nov. Egg mass.

Central nervous system (Fig. 6A)

All of the ganglia are coalesced into a circum-oesophageal nerve ring. The
cerebral and pleural ganglia are entirely fused. Extending anteriorly from the
cerebro-pleural ganglia are the rhinophoral ganglia, each of which gives rise to
three nerves. The eyes are situated at the postero-lateral base of the cerebro-
pleural ganglia. The pedal ganglia are joined by a pair of elongate commissures.
The paired statocysts are situated at the antero-medial ends of the pedal ganglia.

Reproductive system (Fig. 6B)

The ovotestis consists of numerous lobes, each with separate male and
female acini. The preampullary duct is short and expands abruptly into the coiled
ampulla. The ampulla divides into a short oviduct that enters the albumen gland
and the vas deferens. The albumen gland is small and ovoid. The adjacent
membrane gland consists of two lobes. The largest portion of the female gland
mass is the mucous gland. The spherical receptaculum seminis joins the oviduct
near its entrance into the albumen gland by means of a narrow, elongate duct.
The vas deferens expands into a prostatic portion that traverses the dorsal surface
of the penis. From there it extends ventrally and ultimately enters the ental
portion of the penis. The penial papilla is emarginate at its apex.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 245

Development

The egg mass (Fig. 10) is yellowish in life and consists of up to four complete
whorls. There is a single egg per capsule, which develops into a veliger larva with
a Type 1 larval shell (Thompson 1961).

Natural history

All specimens in this study were found feeding upon or in direct association
with the zoanthid anthozoan Palythoa nelliae. Egg masses were commonly
observed upon the zoanthid polyps. Specimens were found near the bases of the
densely arranged polyps during the day or were observed to feed upon the
extended tentacles at night. On several occasions, egg masses of Baeolidia
palythoae were observed to be preyed upon by the aeolid nudibranch Favorinus
Japonicus Baba, 1949.

Berghia chaka sp. nov.
Figs 1C, 11-14

Type material

Holotype. SAM—A35634: Jesser Point, Sodwana Bay National Park, Natal;
intertidal zone; 6 May 1982; collected by T. M. Gosliner.

Paratype. SAM—A35633: Jesser Point, Sodwana Bay National Park, Natal;
intertidal zone; 8 May 1982; collected by T. M. Gosliner.

Etymology

Chaka is the name of a Zulu chief.

Distribution

This species is known only from the type locality, Jesser Point, Sodwana Bay
National Park.

External morphology

The living animals (Fig. 1C) reach 10 mm in length. The oral tentacles are
slender and tapered. The rhinophores (Fig. 11A) are approximately equal to
the oral tentacles in length and possess scattered, elongate papillae. The foot
(Fig. 11B) is moderately broad. Anteriorly it is deeply incised and slightly more
posteriorly a transverse groove is present. The anterior end of the foot is broad
and rounded, without angular or tentacular extensions. The cerata (Fig. 11C) are
irregularly shaped with a few tubercles along their lengths. The anterior right
digestive branch consists of a single arch of eight cerata (Fig. 11D, E). The first
branch of the right posterior digestive system is also an arch composed of ten
cerata. The two to three branches posterior to this may consist of partial arches or
linear rows of two to six cerata. The gonopores are situated ventral to the first
ceratal arch while the anus 1s located within the second arch. The nephroproct is
immediately anterior to the second ceratal arch.

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sr

»)
So

A

Fig. 11. Berghia chaka sp. nov. A. Rhinophore. Scale=1,0 mm. B. Ventral
view. Scale=1,0 mm. C. Ceras. Scale =0,25 mm. D. Lateral view of holotype.
Scale=1,0 mm. E. Lateral view of paratype. Scale = 1,0 mm.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 247

Coloration

The living animals were translucent white covered with a dense pattern of
Opaque white pigment over much of the body. The head and basal halves of the
rhinophores and oral tentacles are covered with chocolate-brown pigment. Brown
is also present at the bases of the cerata and more diffusely on the dorsal portion
of the foot. The digestive gland within the cerata is rusty brown. The cerata are
ornamented with opaque white pigment, which is most dense on their anterior
side and on the irregular tubercles. The tips of the cerata are opaque white. A thin
subapical band of chocolate brown Is present, as is a wider band of opaque white.

Digestive system

A single pair of ovoid oral glands is present along the sides of the buccal mass
(Fig. 12A). They are approximately two-thirds the length of the buccal mass. The

Fig. 12. Berghia chaka sp. nov. A. Buccal region. Scale = 0,5 mm.
B. Jaw. Scale=0,5 mm. C. Reproductive system. Scale = 0,25 mm.

248

ANNALS OF THE SOUTH AFRICAN MUSEUM

seseneenennongonannntnneannennsanntnn ns innannansesansnrgmanens a Sih peraoeNer esas nen -

393X {SKU WO:29mM ¢ 90024

Fig. 13. Berghia chaka sp. nov. Scanning electron micrographs of radular teeth.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 249

Fig. 14. Berghia chaka sp. nov. Scanning electron micrographs of radular teeth.

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

jaws (Fig. 12B) are ovoid with an elongate, smooth masticatory border. The
radula (Figs 13, 14) contains 11 deeply emarginate teeth with 34—36 denticles on
either side of the linear or triangular central cusp.

Reproductive system (Fig. 12C)

The ovotestis is composed of several distinct lobes, each of which contains
both male and female gametes. The ampulla is thin and narrow throughout most
of its length. It is slightly expanded nearest the preampullary duct. The
receptaculum seminis is ovoid and is connected to the ampulla at its bifurcation
into the oviduct and vas deferens. The oviduct is short and enters the yellowish
albumen gland. The membrane gland is slightly smaller than the albumen gland.
The mucous gland is large and lobate. The vas deferens is elongate and expands
slightly into a short prostatic section, terminating at the simply rounded penial

papilla.

DISCUSSION
GENERIC SUBDIVISION

The Aeolidiidae differ from other aeolidacean nudibranchs in that they
possess pectinate rather than cuspidate radular teeth. Within the family generic
relationships have long been controversial (Marcus 1958; Haefelfinger & Stamm
1959; Tardy 1962; Burn 1969; Edmunds 1969; Marcus & Marcus 1970; Gosliner
1980; Gosliner & Griffiths 1981; Rudman 1982). This taxonomic confusion
revolves around the fact that generic boundaries have been based on several
characters that produce conflicting and often polyphyletic taxa. In some cases,
incomplete or erroneous descriptions have compounded the problem. Most
genera have been based on the type of branching of the digestive system within
the cerata or on the degree of elaboration of the primary chemosensory organs,
the rhinophores.

The rhinophores may be papillate (Berghia, Baeolidia), perfoliate (Spurilla
neapolitana), ornamented with ribs (Spurilla macleayi), with bulbous swellings
(Aeolidiella alba, A. japonica), or simple in the remaining members of the family.
Simple rhinophores probably represent the most primitive configuration in the
majority of aeolidaceans. Functionally, increased ornamentation increases the
surface available for sensory detection.

The most primitive form of ceratal arrangement in the Aeolidacea consists of
numerous irregularly spaced rows, which are congested along the edges of the
notum. Within the Aeolidiidae this configuration appears to be present in
Pleurolidia and Protaeolidia. In other members of the family the cerata may be
arranged in linear rows or horseshoe-shaped arches. In most aeolids the digestive
system is divided into anterior and posterior branches. Within the Aeolidiidae
there are also representatives within several genera that have several well-defined
rows in the anterior branch of the digestive system. From this arrangement it is
possible to derive both a configuration with a few rows and one with a single

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 251

anterior arch. Aeolidiella chromosoma possesses five to six rows in the anterior
digestive branch and horseshoe-shaped arches in the posterior branches (Marcus
1961; present study).

Many other aspects of the morphology of the Aeolidiidae vary considerably
and should be discussed within the context of their occurrence within the family.
In the more primitive aeolidaceans, Notaeolidia and the Flabellinidae, the anus 1s
situated ventral to the cerata and is considered to be pleuroproctic (Odhner
1939). Within the Aeolidiidae, this anal position is present in Pleurolidia juliae,
Protaeolidia atra and some members of the genus Cerberilla. It is interesting to
note that Pleurolidia juliae retains another feature characteristic of more
primitive aeolidaceans. It is the only member of the Aeolidiidae that possesses a
lateral tooth on either side of the rachidian tooth (Burn 1966) as in Notaeolidia,
Flabellinidae and Eubranchidae. In Aeolidiopsis ransoni the anus is situated
dorsal to the cerata in the acleloproctic position, as in the Eubranchidae and
Tergipedidae (Pruvot-Fol 1956; Rudman 1982). In the remainder of the
Aeolidiidae the anus is cleioproctic and is located within the ceratal rows or
arches.

The shape of the radular teeth, evenly curved versus emarginate, varies
considerably in the Aeolidiidae (Figs 15, 16) and may vary intraspecifically
(Marcus 1955; Gosliner 1980). In several species of aeolidiids, Berghia major
(Fig. 16B), B. norvegica, Spurilla neapolitana, Baeolidia moebii and B. benteva,
the radula is strongly tapered with the newest teeth being up to seven times the
width of the oldest. In other species, Aeolidiella alba, A. indica (Fig. 15A),
A. chromosoma (Fig. 15B), Berghia chaka and Baeolidia palythoae, the teeth are
uniform in width or increase only slightly.

The masticatory border of the jaws is smooth in most species but may be
denticulate in Berghia major (Gosliner 1980), B. salaamica (Rudman 1982) and
Spurilla neapolitana (Bergh 1877), or papillate in Baeolidia palythoae (present
study). In S. neapolitana the denticulation of the jaws varies intraspecifically.
Within a single population of B. major the jaws may be denticulate or smooth
(Gosliner 1980).

The presence or absence of oral glands in species of the Aeolidiidae was
noted by Rudman (1982). These glands (Table 1) may be shorter than the buccal
mass or may be far more elongate. The glands may consist of a few scattered
vesicles, as in Aeolidiella indica (Fig. 17A), or numerous small or large vesicles.

Rudman (1982: 167) described the reproductive system of Aeolidiopsis
ransoni and stated ‘typical of the family’. However, he described the presence of a
distal bursa copulatrix rather than a proximal receptaculum seminis, a condition
that has not previously been described in any other member of the family. He also
stated that the reproductive systems of Berghia major, B. australis and
B. salaamica were identical to that described for Aeolidiopsis ransoni. This is
clearly contradictory to the configuration previously described for B. major
(Gosliner 1980), which has been re-examined and confirmed in the present study
(Fig. 18A). In Protaeolidia atra both a bursa copulatrix and a receptaculum

DSW

ANNALS

OF THE SOUTH AFRICAN MUSEUM

2OKY

yeaneecnnecennecornagpeccnescren:intenaetornantnnee~necnnneenentcnesontesmannees core

‘ennisiin,

Fig. 15. Scanning electron

‘
&

NAAN ANALOG

8

aN

micrographs of radular teeth. A. Aeolidiella indica

Bergh, 1888. B. Aeolidiella chromosoma (Cockerell & Eliot, 1905).

}
|

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

B 400M 20KY 00 oO9 's§

Fig. 16. Scanning electron micrographs of radular teeth. A. Berghia verrucicornis
(Costa, 1867). B. Berghia major Eliot, 1903.

253

254

Protaeolidia atra*
Aeolidiopsis ransoni
Aeolidia papillosa*
Cerberilla bernadettae
C. affinis*
Aeolidiella alba*

A. indica*

A. chromosoma*

A. olivae*

Spurilla neapolitana*
Berghia major*

B. verrucicornis*

- B. australis

B. salaamica

B. chaka*
Baeolidia benteva*
B. harrietae

B. palythoae*

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Oral glands of the Aeolidiidae.

Oral glands

absent
absent
absent
elongate
elongate
elongate
elongate
elongate
elongate
elongate
absent
elongate
short
short
short
elongate
absent

2 pairs

(1 short,
1 elongate)

* examined in this study

Fig. 17. A. Aeolidiella indica Bergh, 1888. Buccal region. Scale = 1,0 mm.

Vesicles

9

large, uniform
large, uniform
large, scattered
large, scattered
large, scattered
small, uniform

small, uniform
2,
?
small, uniform
small, uniform

small, uniform

B

B. Aeolidiella

chromosoma (Cockerell & Eliot, 1905). Rhinophore. Scale = 1,0 mm.

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

Ly

<tr

Fig. 18. Reproductive systems. A. Berghia major Eliot, 1903. B. Aeolidiella
indica Bergh, 1888. C. Berghia verrucicornis (Costa, 1867). D. Cerberilla affinis
Bergh, 1888.

Nn

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

seminis appear to be present (present study). This configuration is also present in
Aeolidiella indica (Fig. 18B), but the bursa is greatly reduced in size.

In Berghia major a distinct penial gland is present near the gonopore
(Fig. 18A). The only other species of Aeolidiidae in which a penial gland has been
reported is Baeolidia nodosa (Schmekel 1970).

Schmekel (1970) showed that the duct of the receptaculum seminis of
Baeolidia nodosa is wider in the portion nearest the receptaculum and then
sharply diminishes in the portion nearest the hermaphroditic duct. Gosliner
(1980) erroneously reported that the receptaculum of B. nodosa is bilobed. Re-
examination of the single specimen shows that the wider portion of the
receptaculum was mistaken for a second lobe and that the configuration is
actually identical to that described by Schmekel.

Opisthobranch molluscs exhibit a high degree of parallel evolution (Ghiselin
_ 1966; Gosliner 1981; Gosliner & Ghiselin 1984). The Aeolidiidae are certainly no
exception. The incompatability of the subdivision of the family into genera by
means of rhinophores and digestive system attests to the presence of parallelism.
The question remains as to which attributes should be employed to produce a
‘natural’ classification of taxa. Has the ceratal arrangement evolved in parallel or
have the rhinophores, or both? A necessary initial step is to look at other aeolid
families to ascertain which features may have evolved in parallel in other taxa and
which may be unique to the Aeolidiidae. The same evolutionary trends of ceratal
arrangement appear to be present in the closely allied family Facelinidae. In this
taxon, which is likely the sister group of the Aeolidiidae, there are representatives
with numerous rows in the anterior digestive branch as well as more modified forms
with a reduced number of rows or a single arch. Similarly, simple, perfoliate,
knobbed and papillate rhinophores are also present in species of facelinids. Both
ceratal arrangement and rhinophoral elaboration have undergone similar,
parallel evolution between the Aeolidiidae and Facelinidae and therefore provide
little information for the detection of parallelism within the Aeolididae.

Traditionally, possession of ornamented rhinophores has served as the basis
for separating Spurilla from Aeolidiella (Marcus 1961). However, this results in
the combination of species with anterior digestive branch arranged in linear rows
and arches in Spurilla. The ornamented rhinophores within species of Spurilla are
varied in form and include species with knobbed swellings (S. alba, S. japonica),
plicae (S. macleayi), or perfoliations (S. neapolitana, S. olivae, S. chromosoma).
There is little reason to suspect that these ornamentations are monophyletic.
When one examines the species with perfoliate rhinophores there are some
apparent differences in rhinophoral structure. In S. olivae and S. chromosoma
(Fig. 17B) the lamellae are few in number and are arranged diagonally, while in
S. neapolitana there are numerous transverse lamellae. In S. olivae and
S. chromosoma the oral glands contain large isolated glandular bodies as in
Aeolidiella indica (Fig. 17A). In S. alba the glandular bodies are large and evenly
distributed, while in S. neapolitana they are small and uniformly glandular. These
differences strongly suggest that ornamented rhinophores and, in particular, the

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 2.

perfoliate condition has evolved independently within the Aeolidiidae. An
accessory branch of the digestive system (Burn 1969) may have evolved
independently, as well. Gosliner & Griffiths (1981) suggested that perhaps
Spurilla and Aeolidiella could be separated on the basis of ceratal branching as in
Berghia and Baeolidia. The presence of oral glands with large vesicles in species
with the cerata of the anterior digestive branch arranged in rows strengthens this
possibility. Therefore Aeolidiella olivae MacFarland, 1966, Spurilla chromosoma
Cockerell & Eliot, 1905, A. alba Risbec, 1928, and A. japonica Eliot, 1913, are
here regarded as members of Aeolidiella. The form of the rhinophores of
A. orientalis Bergh, 1905, is uncertain and its status must remain open to question.

Rudman (1982) stated that Berghia Trinchese, 1877, should be regarded as a
junior synonym of Spurilla Bergh, 1864, as members of these genera differ only in
the ornamentation of the rhinophores (papillate in Berghia and perfoliate in
Spurilla). The same argument can be applied to Berghia and Baeolidia as they
differ only in the branching of the cerata into rows or arches. The important
question is not whether differences are significant enough to warrant generic
separation, but whether ceratal arches (Berghia and Spurilla) or papillate
rhinophores (Berghia and Baeolidia) are monophyletic in these three taxa. This
question cannot be definitively answered at present. Until such time that it can be
answered, it is preferable to retain the genera as distinct. Spurilla australis
Rudman, 1982, and S. salaamica Rudman, 1982, are transferred to Berghia,
based on the fact that they possess papillate cerata and that the anterior digestive
branch forms an arch.

Rudman (1982) described Aeolidiopsis harrietae and compared it to
Aeolidiopsis ransoni and other taxa with papillate cerata. He stated (p. 160) that
‘placing this species in a genus presents some difficulties’. The discovery of
Baeolidia palythoae in this study places Rudman’s discussion in a different light.
Baeolidia palythoae, A. ransoni and A. harrietae feed exclusively on zoanthid
anthozoans. If one examines aspects of their morphology (Table 2) one can see
that A. harrietae is more similar to B. palythoae than to A. ransoni. Rudman
noted that A. ransoni and A. harrietae are unique among described aeolidiids in
possessing an anterior flange of the jaw but questioned whether this may be a
result of convergence. Both species lack oral glands, but this is also true of
Aeolidia papillosa (present study) and Berghia major (Rudman 1982; present
study). Rudman also noted that, in contrast to other members of the Aeolidiidae,
A. ransoni and A. harrietae have a rounded anterior end of the foot. This is also
true of Baeolidia palythoae and Berghia chaka (present study). Aeolidiopsis
ransoni has radular teeth that are similar in form to those of Protaeolidia (Baba
1955) and Pleurolidia (Burn 1966). Aeolidiopsis ransoni is unique among
described members of the family in possessing an acleioproctic anus dorsal to the
notal brim. Based on these facts A. harrietae more closely resembles species of
Baeolidia, particularly B. palythoae. It is therefore transferred to Baeolidia.

Despite difficulties in separating genera within the Aeolidiidae, it is possible
to differentiate between them on the basis of existing morphological data.

ANNALS OF THE SOUTH AFRICAN MUSEUM

258

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AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 259

KEY TO THE GENERA OF THE AEOLIDIIDAE

Pe MSI ICUTOPLOCHC. oo re Bene ee ene es 2 a erieme, Se once asa ne 2
PMUUSEAC 1OPIOCUE Ol ClElOPNOCUC = = 4k 505) ee ee ee eee -.
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epimoOprockpreanal . 22 o5 5 ley a+ eer Hocus waned weeks ho Sees Aeolidiella

Based on the taxonomic changes proposed here the genera Aeolidiella,
Baeolidia, Berghia and Spurilla include the following species:

Aeolidiella Bergh, 1867

Type species. Aeolidiella soemmeringi (Leuckart, 1828) Bergh, 1867, non
Leuckart = A. alderi (Cocks, 1852), by subsequent designation (Suter 1913).

1. Aeolidiella alba Risbec, 1928

Aeolidiella alba Risbec, 1928: 261, fig. 87, pl. 10 (fig. 9).
Spurilla alba (Risbec, 1928) Edmunds, 1969: 465, fig. 9.

2. Aeolidiella alderi (Cocks, 1852)

Bolisvaldert Cocks, 1852: 1, pl. I (hig. 1).

Aeolidiella alderi (Cocks, 1852) Bergh, 1867: 99.

Aeolidiella soemmeringi (Leuckart, 1828) Bergh, 1867: 99, non Leuckart. Gosliner & Griffiths,
HOSS 121.

3. Aeolidiella chromosoma (Cockerell & Eliot, 1905) comb. nov.
Spurilla chromosoma Cockerell & Eliot, 1905: 51.

4. Aeolidiella drusilla Bergh, 1900
Aeolidiella drusilla Bergh, 1900: 233, pl. 20 (figs 41-46).

5. Aeolidiella faustina Bergh, 1900
Aeolidiella faustina Bergh, 1900: 235, pl. 20 (figs 39, 40).

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

6. Aeolidiella glauca (Alder & Hancock, 1845)

Eolis glauca Alder & Hancock, 1845: 314.
Aeolidiella glauca (Alder & Hancock, 1845) Bergh, 1888: 781.

7. Aeolidiella indica Bergh, 1888
Aeolidiella indica Bergh, 1888: 755, pl. 78 (figs 1, 2).

For a full synonymy see Gosliner & Griffiths (1981: 119).

8. Aeolidiella japonica Eliot, 1913

Aeolidiella japonica Eliot, 1913: 43.
Spurilla japonica (Eliot, 1913) Burn, 1969: 98.

9. Aeolidiella occidentalis Bergh, 1874
Aeolidiella occidentalis Bergh, 1874: 397, pl. 8 (figs 9-19).

10. Aeolidiella olivae MacFarland, 1966

Aeolidiella olivae MacFarland, 1966: 373, pl. 62 (figs 4-6), pl. 72 (figs 9-14).
Spurilla olivae (MacFarland, 1966) Sphon & Lance, 1968: 81.

11. Aeolidiella risbeci (Marcus, 1961)

Aeolidiella ?takanosimemsis (non Baba, 1930) Risbec, 1956: 31, figs 110-115.
Spurilla risbeci Marcus, 1961: 56.

12. Aeolidiella sanguinea (Norman, 1877)

Eolis sanguinea Norman, 1877: 517.
Aeolidiella sanguinea (Norman, 1877) Bergh, 1888: 755.

. Baeolidia Bergh, 1888
Type species. Baeolidia moebii Bergh, 1888: 777, by monotypy.

1. Baeolidia benteva Marcus, 1958
Baeolidia benteva Marcus, 1958: 65, figs 105-111.

2. Baeolidia cryoporos Bouchet, 1977
Baeolidia cryoporos Bouchet, 1977: 60, figs 26, 27.

3. Baeolidia fusiformis Baba, 1949
Baeolidia fusiformis Baba, 1949: 113, figs 158, 159, pl. 50 (fig. 169).

4. Baeolidia harrietae (Rudman, 1982) comb. nov.
Aeolidiopsis harrietae Rudman, 1982: 157, figs 3C, 5D-F, 7-9.

5. Baeolidia moebii Bergh, 1888
Baeolidia moebii Bergh, 1888: 778, pl. 79 (figs 10-16), pl. 80 (figs 1-4).

AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

6. Baeolidia nodosa (Haefelfinger & Stamm, 1959)

Limenandra nodosa Haefelfinger & Stamm, 1959: 420, fig. 1.
Baeolidia nodosa (Haefelfinger & Stamm, 1959) Gosliner, 1980: 66, fig. 19.

7. Baeolidia palythoae sp. nov.

Berghia Trinchese, 1877

Type species. Berghia coerulescens (Laurillard, 1830), by monotypy.

1. Berghia australis (Rudman, 1982) comb. nov.

Spurilla australis Rudman, 1982: 164, figs 12-16, 20.
2. Berghia chaka sp. nov.

3. Berghia coerulescens (Laurillard, 1830)
See Tardy (1962) for synonymy.

4. Berghia creutzbergi Marcus & Marcus, 1970
Berghia creutzbergi Marcus & Marcus, 1970: 87, figs 145-147.

5. Berghia dela Marcus & Marcus, 1960
Berghia dela Marcus & Marcus, 1960: 924, figs 83-86.

6. Berghia japonica (Baba, 1933)

Baeolidia japonica Baba, 1933: 282, fig. 8.
Berghia japonica (Baba, 1933) Marcus, 1958: 68.

7. Berghia major Eliot, 1903
See Gosliner (1980) for synonymy.

8. Berghia norvegica Odhner, 1939
Berghia norvegica Odhner, 1939: 85, figs 52-59.

9. Berghia salaamica (Rudman, 1982) comb. nov.

Spurilla salaamica Rudman, 1982: 173, figs 21A—C, 22, 23.

10. Berghia verrucicornis (Costa, 1867)

Flabellina verrucicornis Costa, 1867: 35, pl. 2 (fig. 4).
Berghia verrucicornis (Costa, 1867) Tardy, 1962: 4.

261

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Spurilla Bergh, 1864
Type species. Eolis neapolitana Delle Chiaje, 1823, by monotypy.

1. Spurilla macleayi (Angas, 1864)
Aeolis macleayi Angas, 1864: 65, pl. 6 (fig. 4).
Spurilla macleayi (Angas, 1864) Burn, 1969: 96, figs 46-50.

2. Spurilla neapolitana (Delle Chiaje, 1823)
See Gosliner (1980) for synonymy.

DISCUSSION OF BAEOLIDIA PALYTHOAE SP. NOV.

Baeolidia palythoae closely resembles B. harrietae (Rudman, 1982) in much
of its anatomy (Table 2). These two species are the only members of the genus
with a single row of cerata in the anterior digestive branch and an anteriorly
rounded foot (Table 3). Although the two species are similar in external
appearance, they differ in several significant features. Baeolidia palythoae
possesses a dark subapical ceratal gland, which is absent in B. harrietae. In
B. harrietae there are four to nine cerata per row, while in B. palythoae there are
never more than five cerata per row. In B. harrietae the gonopore is located at the
anterior end of the first ceratal row, while in B. palythoae it is situated below the
middle of the first row. Baeolidia palythoae is characterized by two distinct pairs
of oral glands, an elongate pair, which 1s also present in B. benteva, and a more
anterior ovoid pair, which exits on the ventral side of the head, by means of
prominent pores. Baeolidia harrietae lacks oral glands. The jaws have a
denticulate margin in B. harrietae, while the masticatory border is papillate in
B. palythoae. In B. harrietae the radular teeth have a maximum of 26 denticles per
side, while in B. palythoae there are 41-55 denticles per side.

DISCUSSION OF BERGHIA CHAKA SP. NOV.

Berghia chaka differs from all previously described species in possessing an
anteriorly rounded foot, rather than one with tentacular foot corners (Table 4). In
external appearance B. chaka most closely resembles B. japonica (Baba, 1933)
(Baba 1949: pl. 50 (figs 168, 169)), which, however, possesses tentacular foot
corners. The jaws in both specimens of B. chaka have a smooth masticatory
border, as do the majority of members of the genus. The degree of emargination
of the radular teeth varies intraspecifically in some species of Berghia, but in
B. chaka all the teeth are deeply emarginate. In some members of the genus the
radular teeth increase markedly in width from the oldest to newest teeth, but in
B. chaka all of the teeth are of about the same width. Oral glands are present in
most species of Berghia, but are absent in B. major (Rudman 1982; present
study). The glands are shorter than the buccal mass in the remaining species,
except in B. verrucicornis and B. coerulescens, where they are much longer than
the buccal mass. The reproductive system of B. chaka contains a proximal
receptaculum seminis as described in B. norvegica (Odhner 1939), B. coerulescens
(Tardy 1962) and B. major (Gosliner 1980).

263

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA 265

ACKNOWLEDGEMENTS

I would like to thank the many people who assisted in the collection of
specimens. Rudiger Bieler first collected Baeolidia palythoae and brought the
species to my attention. Bill Liltved, Mark Cooke, Bonnie Gosliner, and Michelle
and Schalk van der Merwe all collected additional material of Baeolidia
palythoae. Victor Krantz printed some of the final photographic prints and his
help was greatly appreciated.

REFERENCES

ALDER, J. & Hancock, A. 1845. Notice of a new genus and several new species of
nudibranchiate Mollusca. Ann. Mag. nat. Hist. (2) 16: 311-316.

ANGAS, G. 1864. Description d’espéces nouvelles appartenant a plusieurs genres de mollusques
nudibranches des environs de Port-Jackson (Nouvelle-Galles du Sud), accompagnée de
dessins faits d’apres nature. J. Conch., Paris (3) 12: 43-70.

Basa, K. 1930. Studies on Japanese nudibranchs. 3. Venus, Kyoto 2: 117-125.

BaBa, K. 1933. Supplementary note on the Nudibranchia collected in the vicinity of the
Amakusa Marine Biological Laboratory. Annotnes zool. jap. 14: 273-283.

Basa, K. 1949. Opisthobranchia of Sagami Bay collected by his Majesty the Emperor of Japan.
Tokyo: Iwanami shoten.

Basa, K. 1955. Opisthobranchia of Sagami Bay. Supplement. Tokyo: Iwanami shoten.

BARNARD, K. 1927. South African nudibranch Mollusca, with descriptions of new species, and a
note on some specimens from Tristan d’Acunha. Ann. S. Afr. Mus. 25: 171-215.

BERGH, R. 1864. Anatomiske Bidrag til Kundskab om aeolidierne. K. danske Vidensk. Selsk.
Sic (©) 72 139-316.

BERGH, R. 1867. Phidiana lynceus og Ismalia monstrosa. Vidensk. Meddr dansk naturh. Foren.
7-9: 97-130.

BERGH, R. 1874. Beitrage zur Kenntniss der Aeolidiaden. 2. Verh. zool.-bot. Ges. Wien 24:
395-416.

BerGH, R. 1877. Beitrage zur Kenntniss der Aeolidiaden. 5. Verh. zool.-bot. Ges. Wien 27:
807-840.

BerGH, R. 1888. Nudibranchien vom Meere der Insel Mauritius. Jn: Semper, C. Reisen im
Archipel der Philippinen. II. Wiss. Result. ti. Malacol. Untersuch. 16 (1): 755-814.

BerGcH, R. 1900. Ergebnisse einer Reise nach dem Pacific (Schaunisland 1896-1897). Die
Opisthobranchier. Zool. Jb. (Syst. Okol. Geog. Tiere) 13: 207-246.

BeErGH, R. 1905. Die Opisthobranchiata der Siboga Expedition. Siboga Exped. 50: 1-248.

BerGH, R. 1907. The Opisthobranchiata of South Africa. Trans. S. Afr. phil. Soc. 17: 1-144.

Boucuet, P. 1977. Opisthobranches de profondeur de l’océan Atlantique. II. Notaspidea et
Nudibranchiata. J. molluscan Stud. 43: 28-66.

Burn, R. 1966. Descriptions of Australian Eolidacea (Mollusca—Opisthobranchia). 4. The
genera Pleurolidia, Fiona, Learchis and Cerberilla from Lord Howe Island. J. malac. Soc.
Aust. 10: 21-34.

Burn, R. 1969. A memorial report on the Tom Crawford collection of Victorian Opisthobran-
chia. J. malac. Soc. Aust. 12: 64-106.

COCKERELL, T. & Extot, C. 1905. Notes on a collection of Californian nudibranchs. J. Malac.
12: 31-53.

Cocks, W. 1852. New species of Mollusca. Naturalist (Morris) 2: 1.

Costa, A. 1867. Sui molluschi eolididei del Golfo di Napoli. Annuar. Mus. zool. Univ. Napoli 4:
26-37.

DELLE Cuts, S. 1823. Memorie sulla storia e notomia degli animali senza vertebre del regno di
Napoli. Naples.

Epmunps, M. 1969. Opisthobranchiate Mollusca from Tanzania. I. Eolidacea (Eubranchidae
and Aeolidiidae). Proc. malac. Soc. Lond. 38: 451-469.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Epmunps, M. & Just, H. 1983. Eolid nudibranchiate Mollusca from Barbados. J. molluscan
Stud. 49: 179-184.

Exror, C. 1903. On some nudibranchs from East Africa and Zanzibar. 2. Proc. zool. Soc. Lond.
1903: 250-257.

Eviot, C. 1913. Japanese nudibranchs. J. Coll. Sci. imp. Univ. Tokyo 35: 1-47.

GHISELIN, M. 1966. Reproductive function and the phylogeny of opisthobranch gastropods.
Malacologia 3: 327-378.

GosLINER, T. 1980. The systematics of the Aeolidacea (Nudibranchia: Mollusca) of the
Hawaiian Islands with the descriptions of two new species. Pacif. Sci. 33: 37-77.

GosLINER, T. 1981. Origins and relationships of primitive members of the Opisthobranchia
(Mollusca: Gastropoda). Biol. J. Linn. Soc. 16: 197-225.

GOSLINER, T. & GHISELIN, M. 1984. Parallel evolution in opisthobranch gastropods and its
implications for phylogenetic methodology. Syst. Zool. 33: 255-274.

GOSLINER, T. & GrirFiTHs, R. 1981. Description and revision of some South African
aeolidacean Nudibranchia (Mollusca, Gastropoda). Ann. S. Afr. Mus. 84: 105-150.

HAEFELFINGER, H. & Stamm, R. 1959. Limenandra nodosa gen. et spec. nov. (Nudibranchia,
Aeolidiidae prop.), un opisthobranche nouveau de la Méditerranée. Vie Milieu 9: 418-423.

LAURILLARD, C. 1830. In: Cuvier, G. Régne animal 3. Paris: Deterville.

LeuckarT, F. 1828. Breves animalum quorundum maxima ex parte marinorum descriptiones.
Heidelberg.

MACFaRLAND, F. 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North
America. Mem. Calif. Acad. Sci. 6: 1-546.

Macnae, W. 1954. On some eolidacean nudibranchiate molluscs from South Africa. Ann. Natal
Mus. 13: 1-50.

Marcus, E. 1955. Opisthobranchia from Brazil. Bolm Fac. Filos. Ciénc. Univ. S Paulo (Zool.)
20: 89-261.

Marcus, E. 1958. On western Atlantic opisthobranchiate gastropods. Am. Mus. Novit. 1906:
1-82.

Marcus, E. 1961. Opisthobranch mollusks from California. Veliger 3 (suppl.): 1-85.

Marcus, E. & Marcus, E. 1960. Opisthobranchia aus dem roten Meer und von den Malediven.
Abh. math.-naturw. Kl. Akad. Wiss. Mainz 1959: 871-934.

Marcus, E. & Marcus, E. 1970. Opisthobranchs from Curacao and faunistically related
regions. Stud. Fauna Curacao 33: 1-129.

Norman, A. 1877. On two new British nudibranchiate Mollusca. Ann. Mag. nat. Hist. (4) 20:
517-519.

ODHNER, N. 1939. Opisthobranchiate Mollusca from the western and northern coasts of Norway.
K. norske Vidensk. Selsk. Skr. 1939 (1): 1-93.

Pruvot-FoL, A. 1956. Un aeolidien nouveau des mers tropicales: Aeolidiopsis ransoni n.g.,
n.sp. Bull. Mus. natn. Hist. nat. Paris (2) 28: 228-231.

RisBeEc, J. 1928. Contribution a l’étude de nudibranches Néo-Calédoniens. Faune Colon. fr. 2:
1-328.
RisBec, J. 1956. Nudibranches du Viet-Nam. Mém. Inst. océanogr. Nhatrang 9: 1-34.
RupMAN, W. 1982. The taxonomy and biology of further aeolidacean and arminacean
nudibranch molluscs with symbiotic zooxanthellae. Zool. J. Linn. Soc. 74: 147-196.
SCHMEKEL, L. 1970. Anatomie der Genitalorgane von Nudibranchiern (Gastropoda Euthy-
neura). Pubbl. Staz. zool. Napoli 38: 120-217.

SPHON, G. 1971. New opisthobranch records from the eastern Pacific. Veliger 13: 368-369.

SpHON, G. 1978. Additional notes on Spurilla alba (Risbec, 1928). (Mollusca: Opisthobranchia).
Veliger 21: 305.

SPHON, G. & Lance, J. 1968. An annotated list of nudibranchs from Santa Barbara County,
California. Proc. Calif. Acad. Sci. 36: 73-84.

Suter, H. 1913. Manual of New Zealand Mollusca. Wellington: John Mackay.

Tarpy, J. 1962. A propos des espéces de Berghia (Gasteropodes Nudibranches) des cotes de
France et leur biologie. Bull. Inst. océanogr. Monaco 59 (1255): 1-20.

TuieELe, J. 1925. Gastropoda der Deutschen Tiefsee-Expedition. 2. Wiss. Ergebn. dt. Tiefsee-
Exped. ‘Valdivia’ 17: 38-382.

Tuompson, T. 1961. The importance of the larval shell in the classification of the Sacoglossa and
the Acoela. (Gastropoda Opisthobranchia). Proc. malac. Soc. Lond. 34: 233-238.

TRINCHESE, S. 1877. Descrizione del genere Berghia, Trinchese. Rc. Sess. Accad. Sci. Ist.
Bologna 1877: 151-153.

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AEOLID NUDIBRANCHS FROM TROPICAL SOUTHERN AFRICA

ABBREVIATIONS
— anus mu — mucous gland
— albumen gland n | — nephroproct
— ampulla oa — oral gland aperture
— bursa copulatrix og — oral gland
— cnidosac p —penis
— cerebro-pleural ganglion pe — pedal ganglion
— genital aperture pg — penial gland
— mouth pr — prostate
— membrane gland rs —receptaculum seminis

267

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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific namé must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
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figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
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In describing new’species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

T. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; ‘. .. in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

TERRENCE M. GOSLINER

THE AEOLID NUDIBRANCH FAMILY AEOLIDIIDAE
(GASTROPODA, OPISTHOBRANCHIA)
FROM TROPICAL SOUTHERN AFRICA

Semen UNG 95 PART 7 JUNE 1985 ISSN 0303-2515

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~ ANNALS

CAPE ‘TOWN

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(a) Author’s name and year of publication given in text, e.g.:

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Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIsCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & RarFFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THEELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

IMNNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 ~~ Band
June 1985 Junie
Part 9 Deel

DESCRIPTION OF A NEW SOUTH AFRICAN
ARMINACEAN AND
THE PROPOSED RE-INSTATEMENT OF
tak GENUS ATrPETCA BERGE
(MOLLUSCA, OPISTHOBRANCHIA)

By
ROBERTA J. GRIFFITHS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

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DESCRIPTION OF A NEW SOUTH AFRICAN ARMINACEAN
AND THE PROPOSED RE-INSTATEMENT OF THE GENUS
ATTHILA BERGH (MOLLUSCA, OPISTHOBRANCHIA)

By

7

ROBERTA J. GRIFFITHS

Department of Zoology and Institute of Oceanography,
University of Cape Town

(With 4 figures and 2 tables)

[MS accepted 4 July 1984]

ABSTRACT

The morphology of a previously undescribed arminacean opisthobranch mollusc found off
the Cape Peninsula coast, South Africa, is described. The new form cannot be accommodated in
any of the existing arminacean families and the new family Lemindidae is proposed here. The
anatomy of the new species, Leminda millecra sp. nov., is compared with representatives of the
arminacean families Charcotiidae, Heterodorididae, and Doridomorphidae. Examination of the
literature has shown the necessity of re-instatement of the genus Atthila Bergh, 1899, previously
synonymized with Heterodoris Verrill & Emerton, 1882.

CONTENTS

PAGE
NRET GY LUNG BLOM Bee cue hs bots atic d OS A Re 269
De CSGhIPUOM arate atts soe cepa dae it ce eee ea Hei eer 270
DISCUS SI@ Mle he Bees 4 co ed ha ae 273
PREKMOWIC CO CTINEMES: 5 cieiicek 2) 2.0 2 Sscare cle elle arid eet 279
IRGHCMENCES ee ee ree tt Pete enemy OREO RD ae Fe 2 th eee 279
AMY HNIC EI OTISt asec cere en ee ae ee ee ae 280

INTRODUCTION

The South African arminaceans described to date belong to the families
Arminidae and Janolidae, probably the most frequently encountered arminacean
families in the literature. Members of the remaining families are not globally
widely distributed and many are rarely found. However, a new species with
morphological features allied to the Heterodorididae, Doridomorphidae and
Charcotiidae has been found to be quite common below 30m depth, and
occasionally extending to 10 m depth, off the Cape Peninsula coast.

All specimens, except for one that was dredged, were collected by divers
with the aid of SCUBA. Specimens were dissected under a light microscope. The
new species is described here and its affinities within the Arminacea are

269

Ann. S. Afr. Mus. 95 (7), 1985: 269-280, 4 figs, 2 tables.

DAG ANNALS OF THE SOUTH AFRICAN MUSEUM

discussed. The latter requires examination of the various descriptions of
Heterodoris robusta Verrill & Emerton, 1882.

Type specimens are deposited in the South African Museum collections and
the other material in the Ecological Survey Collection, Zoology Department,
University of Cape Town.

DESCRIPTION

Family Lemindidae fam. nov.

Diagnosis

Body elevated with a frontal veil and mantle margins expanded and
undulating, held dorsally above the body. The digestive gland ramifies extensively
within the mantle margin. The anus opens lateroposteriorly; the radula is
multiseriate and there is a copulatory bursa.

Leminda gen. nov.

Diagnosis

Body robust and elevated; quadrangular in section. A velum is present.
Mantle margin broadly expanded and undulating, held vertically above the body.
Foot large and well developed. Rhinophores smooth and retractile into low-
rimmed sheaths. Radula broad and multiseriate with simple hook-shaped teeth.
Cladohepatic digestive gland ramifying extensively into the mantle margin. Eyes
absent. Copulatory bursa opens between the male and female reproductive
apertures.

Type species

Leminda millecra.

Etymology

The generic name is derived from my daughter’s name, Melinda.

Leminda millecra sp. nov.
Figs 1-3

Material

Holotype. SAM-—A35791: off Sandy Bay, west coast of Cape Peninsula
(34°02'S 18°19’E); 36 m depth; 3 January 1981; collected by T. M. Gosliner.

Paratypes. SAM—A35792: 3 specimens; off Sandy Bay, west coast of Cape
Peninsula (34°02'S 18°19’E); 36 m depth; 3 January 1981; collected by T. M.
Gosliner.

Other material. 1 specimen; Whittle Rock, False Bay, Cape Province
(34°15'S 18°33'E); 36 m depth; 27 December 1980; collected by W. R. Liltved.

A NEW SOUTH AFRICAN ARMINACEAN OPISTHOBRANCH Pal

4 specimens; Castle Rock, False Bay, Cape Province (34°18'S 18°29’E); 13 m
deities 23ekebruary 1972 and 27 April 1973; collected™ by (Re J2 Giitfiths.
1 specimen; off Mzimhlava River, Transkei (31°32,2’S 29°42,8’E); dredge;
15 August 1981; collected by R. N. Kilburn.

Etymology

The species name is derived from a combination of the names in my
daughter’s maternal ancestry (Imrie, Clark and Leman).

External morphology

The body is large (Fig. 1A), up to 60 mm or more in length, with a soft and
smooth surface. The muscular foot is broad, square in front, tapering posteriorly

Fig. 1. External morphology of Leminda millecra sp. nov. A. Lateral view of whole
animal. B. Dorsal view of head. C. Ventral view of head. D. Vertical section
through rhinophore cavity.

DD ANNALS OF THE SOUTH AFRICAN MUSEUM

to a short blunt tail. A large velum extends anterior to the mouth (Fig. 1B, C).
The mantle edge is enlarged into a continuous broad undulating margin,
orginating between the rhinophores, wide in front and narrowing, but con-
tinuous, over the tail region (width 12 mm to 4 mm respectively in a preserved
60 mm animal). It is held vertically over the dorsum and contains extensively
ramified digestive gland. The rhinophores are smooth and can be retracted into
sheaths that bear a narrow collar (Fig. 1D). Eyes are absent. The positions of the
genital apertures, nephroproct and anus are shown in Figure 1A.

Colour

The body is translucent white with dark brown to black digestive gland
visible in the mantle margin. The epidermis is coloured to varying intensity with
‘luminous’ blue pigment. Blue colouring extends along the edges of the frontal
veil, foot and mantle margin, and the distal third of the rhinophores. In some
specimens extensive blue pigment may cover the sides of the foot, notum and
mantle margin and the whole animal may appear bright ‘luminous’ blue in colour.
In this case the ramifications of the digestive gland appear black, becoming deep
purple towards the edge of the notum. The digestive gland stops several
millimetres short of the edge of the mantle.

Internal morphology

The anterior third of the large muscular buccal mass is covered by jaws
(Fig. 2A), which are heavily chitinized only along the smooth cutting edge
(Fig. 2B). The broad radula bears a variable number of teeth per row. Radulae
from two large specimens showed formulae of 37 X 51-70.1.70-51 and
39 x 73-92.1.92-73, and from a smaller specimen (50 mm _ preserved),
27 X c. 52.1.52. All teeth are simply hamate, the rachidian stouter than the
laterals, and of uniform size, except for the outer laterals, which diminish in size
towards the edge of the radula (Figs 2C, D).

Large oral and salivary glands are present (Fig. 3A), the latter ramifying
extensively over the dorsal surface of the digestive and reproductive systems. A
large crop precedes the stomach, from which two anterior and two posterior
branches of the digestive gland arise (Fig. 3A) and ramify within the mantle
margin. There are no discrete digestive gland lobes. The intestine extends dorso-
laterally over the anterior aorta and opens at the lateral anus in the posterior third
of the body.

The heart lies in the posterior half of the body cavity, dorsal to the intestine
(Fig. 3A). The central nervous system (Fig. 3B) contains cerebro-pleural and
pedal ganglia.

The reproductive system is shown in Figure 3C. The ovotestis fills the
posterior body cavity and a long hermaphrodite duct joins a sac-like convoluted
ampulla, which narrows and branches to the vas deferens. The oviduct is deeply
embedded between the vagina and the female gland masses, and no branch can be
seen before it enters the musculature of the vagina. The mucous gland is

N
|
1S)

A NEW SOUTH AFRICAN ARMINACEAN OPISTHOBRANCH

B
\

3mm
D
Cc C
0,) mm (C

Fig. 2. Leminda millicra sp. nov. A. Lateral view of buccal mass showing
position of jaws. B. Jaws. C. Rhachidian and first two lateral teeth of radula.
D. Sharp unused and blunt used lateral teeth from the radula.

extensive. A receptaculum seminis is absent and the copulatory bursa opens
beside the vaginal aperture. The coiled vas deferens thickens slightly before
entering the muscular penis sac, which bears an elongate conical unarmed penis.

Geographical range

This species has been collected from both east and west coasts of the Cape
Peninsula, and off the Transkei coast of southern Africa. It appears to be a cool-
water form, seldom found at depths less than 30 m.

DISCUSSION

The Arminacea are a heterogeneous group of opisthobranchs distinguished
primarily by the presence of a velum (Odhner 1934) and general absence of
tentacles (except in the genus Goniaeolis and the family Janolidae). Placement of
an individual species within this group is usually based upon the possession of a

DA ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Leminda millecra sp. nov. A. Internal morphology, dorsal view. B. Central nervous system.
C. Reproductive system.

combination of characters rather than any one distinctive feature. In addition to
the presence of a velum, the arminaceans usually possess only a single seminal
vesicle (except in Janolidae), a lateral anus (except in Janolidae), and generally
simple rhinophores without a sheath (Franc 1968). As in the other major
nudibranch groups the Arminacea have representatives with doridiform to
aeolidiform external appearance (Odhner 1934, 1939), a digestive system ranging
from more or less holohepatic to cladohepatic with a tendency towards the latter,
rhinophores retractile or non-retractile, and radulae ranging from broad
multiseriate to reduced with 3 teeth per row. Characters placing Leminda millecra

A NEW SOUTH AFRICAN ARMINACEAN OPISTHOBRANCH 2S

within the Arminacea are the presence of a velum, absence of extended
rhinophore sheaths and tentacles, a lateral anus, cladohepatic digestive gland and
a single seminal vesicle.

Division of the order Arminacea into the sub-orders Euarminacea and
Metarminacea (Franc 1968) is again based on the presence of a combination of
characters, which present either doridiform (primitive) or aeolidiform (advanced)
facies respectively (Odhner 1934). However, the characters constituting “primi-
tiveness’ or ‘advancement’ remain in question (Gosliner 1981) and as many
members of the Arminacea show a combination of aeolid and dorid characters (as
defined by Odhner 1934) this division is untenable. Future reorganization within
the group will no doubt be required once the affinities of the families are better
understood.

Leminda millecra does not belong in any existing family and appears to
possess characters found in the doridiform Heterodorididae and Doridomorphi-
dae, and the aeolidiform Charcotiidae. Table 1 lists the morphological features of
genera within these families. The external body form of L. muillecra most closely
resembles that of Telarma antarctica Odhner, 1934 (Charcotiidae). with its
elevated, quadrangular body, much exposed mantle margins bearing ramified
digestive gland, and its smooth rhinophores. However, the internal structure of
T. antarctica differs in that discrete anterior and posterior lobes of the digestive
gland are present (Fig. 4A), the seminal vesicle joins the female gland mass, and
the radula is reduced to 5 teeth per row. The members of the Charcotiidae are all
Antarctic forms with reduced radulae. Heterodoris robusta Verrill & Emerton,
1882 (Heterodorididae), described in Verrill (1882), has an external body form
similar to L. millecra but the mantle margins are only slightly extended; it has an
elevated body and a narrow and thin undulating mantle margin (Odhner 1926).
Like L. millecra it does not possess eyes and has a broad radula with many simple
hooked teeth. These similarities are contrasted by the following differences: the
digestive gland in H. robusta consists of three large discrete lobes filling the body
cavity (Fig. 4B); the gland has a ramified structure but does not enter the mantle
(Odhner 1926); the rhinophores are perfoliate, the mantle has dispersed tubercles
(Bouchet 1977) and the reproductive system differs in the presence of a prostate;
the seminal vesicle opens into the vagina (Odhner 1926; Bouchet 1977); and a
cylindrical rather than a conical penis is present. The members of the family
Doridomorphidae bear little resemblance to L. millecra. Doridomorpha gardineri
Eliot, 1906, has a flattened dorid shape but with a horizontally broadened mantle
edge. The external and internal anatomy of D. gardineri differs from L. millecra
in several respects: the animal appears to be small; the rhinophores are perfoliate;
the jaws are denticulate; the radula is reduced; and the reproductive system is
triaulic. However, the digestive systems in the two species are very similar
(Fig. 4C, D). Thus, while some similarities exist, the considerable differences
between L. millecra and the members of the above three families are considered
sufficient to warrant separate familial status and the family Lemindidae is
therefore proposed. The similarities between the above four families are not

ANNALS OF THE SOUTH AFRICAN MUSEUM

276

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showing the stomach, intestine and digestive gland. Stippled area indicates the
presence of discrete lobes of digestive gland. A. Telarma antarctica (after
Odhner 1934). B. Heterodoris robusta (after Odhner 1926). C. Dorido-
morpha gardineri (after Eliot & Evans 1908). D. Leminda millecra sp. nov.

considered sufficient to support their fusion into a single family. The six species
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no single character is common to more than two or three species. There appears
to be no consistent feature or combination of features to link the species together
and serve in diagnosing a joint family.

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descriptions of Heterodoris robusta were reviewed. Examination of the descrip-
tions by Verrill & Emerton, in Verril (1882), Odhner (1926) and Bouchet (1977),
and of Afthila ingolfiana (Bergh, 1899), which was synonymized with H. robusta

ANNALS OF THE SOUTH AFRICAN MUSEUM

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A NEW SOUTH AFRICAN ARMINACEAN OPISTHOBRANCH 279

by Odhner (1926), indicate that the synonymy was not warranted and that
A. ingolfiana should be regarded as a separate species. Odhner’s (1926)
synonymy was based upon examination of the external surface of what remained
of Bergh’s dissected specimen, with little reference to Bergh’s reasonably
adequate description. There remain essential differences in anatomical structure
that have not been taken into account. Reference to Table 2, which lists the
salient features from the original descriptions of the species, shows the following
differences:

1. Atthila ingolfiana, although a smaller specimen, bears more perfoliations
on the rhinophores and more rows of teeth in the radula, indicating that it is not
merely a less mature specimen of H. robusta.

2. Atthila ingolfiana bears a bilobed edge to the rhinopore cavity. This was
confirmed in Odhner’s examination of the specimen and is not present in any
specimens of H. robusta.

3. In A. ingolfiana the size of the lateral teeth in a radula row initially
increases and then decreases towards the radula edge. In H. robusta the lateral
teeth continue to increase in size towards the edge of the radula, the largest being
at the edge.

4. The median and first lateral teeth in A. ingolfiana are denticulate whereas
the teeth of H. robusta showed no denticulations (Odhner 1926). If the
denticulations had been worn from the older teeth in Odhner’s specimens, they
should have been visible on the newer unused teeth in the radula.

5. From the drawing of the reproductive system, A. ingolfiana does not
appear tO possess a prostatic portion of the vas deferens. This is present in
H. robusta.

The above differences indicate that Bergh’s and Odhner’s specimens were
probably representative of different species. Until new material of Asthila and
Heterodoris can be examined, I consider that the genus and species A/fthila
ingolfiana should be maintained as separate from Heterodoris robusta. The genus
Aithila Bergh, 1899, should thus be re-instated, bearing the following diagnosis
(Bergh 1899): Body elevated as in Tritonia, rhinophores perfoliate with bilobed
edge to the rhinophore cavity. Masticatory border smooth, radula multi-seriate
with denticulated median and first lateral teeth. Digestive gland lobes extensive.
but do not ramify into the body wall.

ACKNOWLEDGEMENTS

Dr E. Marcus and Dr T. M. Gosliner are thanked for fruitful discussions. and
Dr R. N. Kilburn for the loan of material.

REFERENCES

BerGu, R. 1899. Nudibranchiate Gasteropoda. Dan. Ingolf-Exped. 2 (3): 1-49.
Boucuet, P. 1977. Opisthobranches de profundeur de locéan Atlantique: II— Notaspidea et
Nudibranchiata. J. molluscan Stud. 43: 28—66.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eutor, C. N. E. 1906. Nudibranchiata with remarks on the families and genera and description of
a new genus, Doridomorpha. In: GARDINER, J. S. The fauna and geography of the Maldive
and Lacadive archipelagoes 2: 540-573.

Euiot, C. & Evans, T. J. 1908. Doridoeides gardineri: a doridiform cladohepatic nudibranch.
Q. Jl microse. Sci. 52: 279-299.

Franc, A. 1968. Mollusques gastéropodes et scaphopodes. Jn: Grasse, P. ed. Traité de zoologie
5 (3). Paris: Masson et Cie.

GosLInER, T. M. 1981. Origins and relationships of primitive members of the Opisthobranchia
(Mollusca: Gasteropoda). Biol. J. Linn. Soc. 16: 197-227.

OpuHNER, N. H. 1926. Nudibranchs and lamellariids from the Trondhjem Fjord. K. norske
Vidensk. Selsk. Skr. 1926 (2): 1-36.

Opune_er, N. H. 1934. The Nudibranchiata. Br. Antarct. Terra Nova Exped. 1910 (Zool.) 7:
229-310.

OpHNER, N. H. 1939. Opisthobranchiate Mollusca from the western and northern coasts of
Norway. K. norske Vidensk. Selsk. Skr. 1939 (1): 1-93.

THIELE, J. 1912. Die antarktischen Schnecken und Muscheln. Dt. Stidpol.-Exped. (Zool. 5) 13:
183-285.

VayssIERE, M. A. 1906. Diagnoses génériques de mollusques gastéropodes nouveaux rapportés
par l’expédition antarctique du Dr. Charcot. Bull. Mus. Hist. nat., Paris 12: 148.

VERRILL, A. E. 1882. Catalogue of marine Mollusca added to the fauna of the New England
region, during the past ten years. Trans. Conn. Acad. Arts Sci. 5: 548-549.

ABBREVIATIONS

a anus n nephridiopore
adgb anterior digestive gland ) ovotestis

branches oe oesophagus
al albumen gland og oral gland
amp ampulla OV oviduct
at atrium p penis
be bursa copulatrix pdgb posterior digestive gland
G cerebral ganglion branches
cp cerebro-pleural ganglion pe pedal ganglion
dg digestive gland it rhinophore
f foot S stomach
ga genital aperture sg salivary gland
1 intestine Vv vagina
me membrane gland ve velum
mf mantle folds ven ventricle
mu mucous gland vd vas deferens

tv

I}

Oh ov

! ay
1 zi
ont
\
4
2
-
6 ;
, ed
.
ba
5
a “i
i :
ve i

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

‘An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-1SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus...’; ‘. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

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ROBERTA J. GRIFFITHS

DESCRIPTION OF A NEW SOUTH AFRICAN
ARMINACEAN AND THE PROPOSED RE-INSTATEMENT
OF THE GENUS ATTHILA BERGH

(MOLLUSCA, OPISTHOBRANCHIA)

JUNE 1985 | ISSN 0303-2515

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BuLLOuGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THEELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band
June 1985 Junie
Rant 8 Deel

A REVISION OF
THE ORNITHISCHIAN DINOSAUR
KANGNASAURUS COETZEEI HAUGHTON,
WITH A CLASSIFICATION OF
THE ORNITHISCHIA

By
MICHAEL R. COOPER

Cape Town Kaapstad

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are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A REVISION OF THE ORNITHISCHIAN DINOSAUR
KANGNASAURUS COETZEEI HAUGHTON,
WITH A CLASSIFICATION OF THE ORNITHISCHIA

By
MICHAEL R. COOPER

National Museum, Bulawayo, Zimbabwe*

(With 22 figures)

[MS accepted 11 July 1984]

ABSTRACT

The osteology of Kangnasaurus coetzeei Haughton is redescribed on the basis of the
hypodigm, and figured in detail. Long considered an iguanodontid, it is here shown to be most
closely allied to Dryosaurus and is thus transferred to the Hypsilophodontidae. To judge from its
evolutionary grade, the early Cretaceous age assigned to Kangnasaurus seems reasonable. The
unsatisfactory higher taxonomy of the Ornithischia, in particular the Ornithopoda, has resulted
in a cladistical analysis of the group and the introduction of a new classification.

CONTENTS

PAGE
MMO CH OM ees. Lire ce hc 8 CE te ae a ar ee cee 281
OirambinischranwtaxOnomy Aelia. oe oss ec sheng ale eae ee ae 283
SNiSieMianle GESCHIPTOM 1... 24 Pe eae ho ae ee ae 293
SUDAN y ct te he Seen eer ea Sverre ceri Co cn Re SHS
PACKMIONVIC CSE MM CMUS 22.58 sie bias sue ae te tO ee ene eee SS
FINGRGHEIN CES ye APs. ssh Site attest aye ae. Wile othe a eRe pea BilS

INTRODUCTION

The occurrence of dinosaur bones at the base of the Kalahari succession in
the northern Cape (Fig. 1) was reported by Rogers (1915), and the find was
described by Haughton (1915). The material was obtained from a poorly sorted,
immature, coarse clastic unit of colluvial rubble exposed in a well at a depth of
approximately 34m. Although there are preservational differences amongst
some of the bones all the femora are undoubtedly conspecific while most of the
other bones are certainly of ornithopod character. Furthermore, there is no
reason to believe that the femora belong any more certainly with the holotype
tooth than do the highly porous dorsal and caudal vertebrae. These preser-
vational differences are here related to postdepositional differential leaching and
the entire collection of dinosaur bones from this well is treated as the hypodigm of
Kangnasaurus coetzeei Haughton. The writer is of the belief that when dealing

* Present address: Department of Geology, University of Durban-Westville, Private Bag
X54001, Durban 4000.

281

Ann. S. Afr. Mus. 95 (8), 1985: 281-317, 22 figs.

to Vioolsdrif

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

to Karasburg

Keetmanshoop
®

® Springbok

@-Cape Town

L
Goodhouse
me NEEL Kar

to O’kiep

Fig. 1. Locality map with fossil site arrowed.

with such disarticulated remains there is a high degree of probability that all the
bones are from individuals of the same taxon. Since, however, this probability
cannot be quantified, the burden of proof must lie with the dissenter to establish
conclusively that more than one taxon is involved. Thus, Steel’s (1969: 19)
statement (taken almost verbatim from Haughton (1915: 259)) that *. . . the foot
bones and vertebrae appear to come from a different deposit to the type and their
inclusion in this genus is questionable’ 1s here rejected. The preservation of the
foot bones and tibiae are identical to those of the femora.

It is clear from the literature that Kangnasaurus 1s a poorly known genus, yet
the available material allows for a better understanding of the taxon than
provided by Haughton (1915). Consequently, it is the purpose of this paper to

OO

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 28

redescribe and illustrate the available material of K. coetzeei, and to show that its
affinities have been misinterpreted previously.

All the material is housed in the palaeontological collections of the South
African Museum under the catalogue numbers SAM—2731 and SAM-—2732.

ORNITHISCHIAN TAXONOMY

Present classification of the Ornithischia, in particular the Ornithopoda, is
unsatisfactory (Maryanska & Osmolska 1974; Santa Luca 1979; Dodson 1980;
Coombs 1982), while earlier classifications (Thulborn 1971, 1975; Galton 1972)
recognized horizontal ‘grades’ rather than monophyletic lineages. Since the writer
follows Santa Luca (1979) in regarding the presence or absence of an obturator
process to the ischium as of prime taxonomic importance, a new classification of
the Ornithischia is proposed (Fig. 2). This is based upon the following character
suites:

Character suite A

Subclass DINOSAURIA Bakker, 1975

Archosaurs with a wide open iliac acetabulum, prominent 4th trochanter,
mesotarsal ankle and ascending process to the astragalus. Pubis long, thickened
distally.

Character suite B

Cohort CARNOSAURIFORMES new cohort

Dinosaurs retaining the primitive condition of recurved thecodontian
dentition with finely serrated cutting edges.

Character suite C

Cohort ORNITHISCHIFORMES new cohort

Dinosaurs with laterally compressed, leaf-shaped teeth bearing marginal
denticles to the cutting edge. Dentition heterodont with non-recessed, marginal
cheek teeth.

Character suite D

Superorder PACHYPODOSAURIA new superorder

Long-necked ornithischiforms that retain the primitive brachyiliac pelvis.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM
5 : Fabrosauridae
\ ger aniesaumiorines Pr artagm
>
W
ie es 2 _®@ Hadrosauridae
za
oO U A
i <6 Iquanodontidiae
ale one
Nee AN
m nN of "Se camptosaumtiee
= Hypsilophodontidae

Sas Scelidosauridae
©
al Stegosauridae
se (Op)
a0) Se

Acanthopholidae
@ Nodosauridae
= # Pisanosauridae
N
x< @ Heterodontosauridae
# Stenopelixidae

a

—+ — ‘® Pachycephalosauridae

uy
ay
‘s

—\ 2L® Protoceratopsidae
ok,

# Psittacosauridae

—sS
\‘e Ceratopsidae

Fig. 2. Hypothesized relationships amongst the Ornithischia.

Character suite E

Superorder ORNITHISCHIA Seeley, 1888 (nom. transl. herein
ex order Ornithischia)

Herbivorous ornithischiforms with wear facets to the cheek teeth, an
opisthopubic pelvis, a supraorbital element, a predentary bone to the mandible,
an inturned head to the femur, and a pendent 4th trochanter.

Nn

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 28

Character suite F

Order ORNITHOPODA Marsh, 1871 (nom. transl. herein
ex suborder Ornithopoda)

Ornithischians with an obturator process to the ischium, at least 24 pre-
sacrals, a jugal of normal size that lacks a prominent ventral flange, and a promi-
nently flared lesser trochanter separated by a deep cleft from the femoral shaft.

Character suite G

Suborder FABROSAURIA new suborder

Ornithopods retaining the primitive condition of non-recessed, marginal
cheek teeth. Lateral surface of maxilla flat, dentary slender. Jaw articulation in
line with tooth row. Six premaxillary teeth. Cheek teeth lack well-defined occlusal
wear surfaces. Prepubis short.

Superfamily FABROSAUROIDEA new superfamily

Diagnosis as for suborder.

Family NANOSAURIDAE Marsh, 1878

Diagnosis as for suborder. Since Galton (1978) includes Nanosaurus within
the Fabrosauridae, the latter taxon (Galton 1972) is a junior subjective synonym
of the family Nanosauridae.

Character suite H

Suborder HyPsSILOPHODONTIA new suborder

Ornithopods with recessed cheek teeth roofed by an overhanging maxilla and
floored by a massive dentary. Cheek teeth markedly asymmetrical, with
prominent ridging of opposing surfaces. Jaw articulation ventrally offset.
Elevated coronoid process and well-developed retroarticular process. Prepubis
long.

Character suite I

Superfamily HypsILOPHODONTOIDEA new superfamily

Hypsilophodonts retaining the primitive ornithopod characters of a short,
high skull with large orbits, premaxillary teeth, small external nares, moderately
developed antorbital vacuities and dermal armour. Diastema separating pre-
maxillary teeth from those in the maxilla. Horny beak anteriorly. Maxillary teeth
lack median ridge, but medial surface of dentary teeth strongly ridged. Posterior
cervicals and dorsal vertebrae amphicoelous. Scapula short, expanded distally.
Anterior intercondylar groove to femur weakly developed or absent.

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family HypstLoPpHODONTIDAE Dollo, 1882

Diagnosis as for superfamily.

Character suite J

Infraorder IGUANODONTIA Dollo, 1888

Large to very large graviportal hypsilophodonts. Head large, long, laterally
compressed, with elongate snout, vestigial antorbital vacuities, and large external
nares. Premaxilla edentulous, separating maxilla from nasal. Nasals elongate.
Teeth unilaterally enamelled, those of the maxilla with a median ridge. Cervical
and anterior dorsal vertebrae opisthocoelous. Scapula long, straight, slender.
Humerus relatively narrow, with weak deltopectoral crest. Posterior process of
ilium relatively produced and somewhat decurved, with brevis shelf. Pubis and
-ischium decurved. Carpus compact, generally well ossified. Digit II] of manus
with three phalanges. Pedal digit I reduced. Astragalus lacks an ascending
process.

Character suite K

Superfamily CAMPTOSAUROIDEA new superfamily

Primitive iguanodonts retaining a single supraorbital and curved femur with
shallow anterior intercondylar groove. Premaxilla expanded, almost encircling
nares. Quadrate short, curved, inclined. Short ventrolaterally directed basi-
pterygoid processes. Phalangeal formula for manus 2—3-—3-3-2. Prepubis
relatively shallow, postpubis as long as ischium.

Family CAMPTOSAURIDAE Marsh, 1888

Diagnosis as for superfamily.

Character suite L

Superfamily IGUANODONTOIDEA Hay, 1902

Iguanodonts with digit I of the manus reduced; digit II with a ‘hoof-like’
ungual. Prepubis relatively deep; postpubis reduced, shorter than ischium. There
are 5-8 sacral vertebrae. Femur straight, columnar, with very deep anterior
intercondylar groove. Pedal phalangeal formula 0—3—4—5-0.

Character suite M

Family IGUANODONTIDAE Cope, 1896

Iguanodontoids with a low skull, two supraorbital elements and a slender
coronoid process. Phalangeal formula of manus 1—3—3—3-—4, with a spur-like
phalanx to the pollex. There are 5-6 sacral vertebrae.

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 287

Character suite N

Family HADROSAURIDAE Cope, 1896

Specialized aquatic iguanodontoids with expanded premaxillaries forming a
duck-like bill. Premaxillae and nasals frequently extensively modified, sometimes
forming a crest. Supraorbital elements lacking. Maxillae and mandibular rami
with dental battery comprising 45—60 rows of successional teeth. There are 30—34
presacrals (approximately 15 cervicals) and usually 8 sacral vertebrae. [lium with
antitrochanter. Ischium straight. Femur with enclosed anterior intercondylar
groove. Pollex absent; phalangeal formula of manus 0—3-—3-3-3.

Character suite O

Order NEORNITHISCHIA new order

Ornithischians lacking an obturator process to the ischium and with recessed
cheek teeth. Humerus with expanded head and prominent deltopectoral crest.
UlIna with distinct olecranon.

The presence of recessed cheek teeth in the neornithischians is a character
shared with hypsilophodonts. As noted by Galton (1973), cheek pouches merely
reflect advanced adaptation to herbivory and hence may have evolved more than
once, in different herbivorous lines. The fact that they are present in the early
Carnian Pisanosaurus but absent in the Hettangian Fabrosaurus supports a
fundamental dichotomy within the Ornithischia.

Character suite P

Suborder THYREOPHORINA Nopsca, 1915

Large to very large, heavily armoured, quadrupedal neornithischians with
edentulous premaxillae and vestigial to absent antorbital vacuities. The cheek
teeth reflect the primitive condition and are laterally compressed, deeply
denticulate and feebly developed. External nares subterminal and laterally
directed. Skull relatively low and long.

Character suite OQ

Infraorder SCELIDOSAURIA new infraorder

Very primitive thyreophorinids, which are relatively weakly armoured
compared to the remainder of the group, with four sacral vertebrae, reduced
upper temporal fenestrae and ossified axial tendons.

Retains such primitive ornithischiform characters as a supra-acetabular
buttress to the ilium, maxillary teeth that close outside those of the dentary (in
prosauropod fashion), the lack of wear facets to the teeth, two distal tarsals,
moderately long spool-shaped amphicoelous cervical vertebrae, distal condyles to
femur with practically straight lateral margin and expanded medial condyle, limb

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

bones comparatively slender with main elements hollow, tibia shorter than femur,
humerus 70 per cent of femoral length, hind foot functionally tridactylous, the
hallux reduced and digit V apparently lacking, and with a pedal phalangeal
formula of 2—3—4—5—0. The number of presacral vertebrae is uncertain; there are
17 dorsals and at least 6 cervicals; perhaps there were 25 presacrals as in the
prosauropods.

There are still considerable problems surrounding Scelidosaurus and it is
seriously in need of modern revision. One of the major sources of controversy
surrounds the status of the ‘juvenile Scelidosaurus’ (Charig 1972; Galton 1975;
Thulborn 1977), which Thulborn believes to be generically distinct and allied to
Fabrosaurus. It differs from the adult holotype in having a very short prepubis,
which led Romer (1968) and Galton (1975) to conclude that it was a primitive
ankylosaur, while Charig (1976) has commented on its ‘ankylosaur-like’ skull.
_ However, a short prepubis is primitive for the ornithischians since it is also found

in Fabrosaurus and Heterodontosaurus. Thulborn (1977) also pointed to the fact
that the postpubis of the juvenile was as long as the ischium whereas that of the
adult was shorter. It is to be wondered whether these differences are not the
result of allometric growth, with juveniles reflecting the primitive condition (as
has been suggested for the prosauropod Euskelosaurus, cf. Cooper 1981).

Zittel (1932), Romer (1956) and Steel (1969) have all treated Scelidosaurus
as a monotypic subfamily within the Stegosauridae. It is, however, so primitive,
and thus resembling ornithopods, that it should certainly be housed in its own
family.

Thulborn (1977) regarded the juvenile Scelidosaurus as closely allied to
Fabrosaurus, but its short prepubis and the lack of an obturator process suggest it
is at least as close to Heterodontosaurus. In addition, the femoral head of
Scelidosaurus is said to be subglobular, perhaps reflecting poor discrimination
between the femoral head and greater trochanter, as found in Heterodontosaurus.
The two genera also have a brevis shelf to the ilium, while the supraorbital of
Scelidosaurus was said to resemble that of Stegoceras (Coombs 1972).

Family SCELIDOSAURIDAE Cope, 1869

Diagnosis as for infraorder.

Character suite R

Infraorder STEGOSAUROMORPHA new infraorder

Thyreophorinids in which the jugal is small, the quadrate inclined and the
upper temporal fenestrae are reduced to absent. External nares moderately large.
Retroarticular and coronoid processes poorly developed to absent. Neck short,
with abbreviated, disc-like centra. Vertebrae platycoelous to amphiplatyan.
Transverse processes of dorsal vertebrae inclined upwards. Ilium with long
anterior and short posterior process. Limb bones solid, or nearly so. Cnemial

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 289

crest of tibia poorly developed. The 4th trochanter is represented by a rugosity
only. Ungual phalanges of pes ‘hoof-like’.

Character suite S

Superfamily STEGOSAUROIDEA Marsh, 1877 (nom. transl. herein
ex suborder Stegosauria)

Stegosauromorphs with a proportionately very small skull and three
supraorbital elements. Neural arches and spines of posterior dorsals, sacrals and
anterior caudals very tall. Sacral cavity greatly enlarged. Femur columnar with
small lesser trochanter and little distinction between the femoral head and greater
trochanter. About 27 presacrals (10 + 17). Ilium with decurved anterior process.
Humerus less than half femoral length. Ventral end of scapula greatly expanded.
Astragalus and calcaneum may fuse with each other and with epipodials.
Metapodials very short. Phalangeal formula of pes 0—3—3-—3-0. Dermal armour
very prominent, comprising large vertical plates and long spines.

Character suite T

Superfamily ANKYLOSAUROIDEA von Huene, 1914 (nom. correct. Osborn, 1923;
nom. transl. herein ex suborder Ankylosauria)

Stegosauromorphs characterized by the massive development of dermal
armour, with skull partially or largely covered by dermal ossifications. Upper
temporal fenestra closed. Lateral temporal fenestra strongly reduced, slit-like, or
occluded by armour. Orbits small, with overhanging supraorbital region. There
are 6—9 sacrals. Anterior process of ilium out-turned to a marked degree. Pubis
greatly reduced; no prepubis and postpubis rudimentary. Ischium strongly curved
distally. Caudal vertebrae short.

Character suite U

Family ACANTHOPHOLIDAE Nopsca, 1902

Moderately sized, primitive ankylosaurs with thin accessory dermal covering
to the skull and only moderately developed dermal armour.

Character suite V

Family NODOSAURIDAE Marsh, 1890

Large to very large ankylosaurs with a comparatively large skull displaying a
short, rounded snout and broad posterior margin. The jugal is deeply sculptured.
A thick dermal covering is present, comprising separately ossified plates of
varying sizes. Atlas—axis usually fused while the dorsal ribs are frequently co-
ossified with the vertebrae. Anterior process of ilium broad. Acetabulum
sometimes closed. Femur massive. Armour very heavy, encasing the tail, and in
the pelvic region frequently fusing with the ilia, vertebrae and ribs.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Character suite W

Suborder NEORNITHOPODA new suborder

Neornithischians with subcylindrical cheek teeth displaying planar wear
surfaces. Jugal with prominent ventral flange. There are 21-22 presacral
vertebrae.

Character suite X

Infraorder HETERODONTOSAURIA new infraorder

Primitive neornithopods retaining a small skull with large orbits, well-
developed antorbital vacuities, and small external nares. Greater trochanter
poorly distinguished from femoral head. Lesser trochanter small, adpressed to
femoral shaft.

Character suite Y

Family PISANOSAURIDAE Casamiquela, 1967

Very primitive heterodontosaurs with an unfused ankle resembling the
prosauropod condition. Cheek teeth apparently unridged (due to wear, according
to Bonaparte 1976). Coronoid process prominent. Retroarticular process
moderately developed. Femur with prominent posterior intercondylar groove.
Dentary massive, much of it lateral to tooth row.

Character suite Z

Family HETERODONTOSAURIDAE Romer, 1966; Kuhn, 1966

Heterodontosaurs with caniniform processes to the premaxilla and dentary.
Nasals bulbous. Prepubis short, deep; postpubis long. Femur lacking anterior and
posterior intercondylar grooves. Tibia—fibula and astragalus—calcaneum fused to
form functional tibiotarsus and tarsometatarsus respectively. Three distal tarsals.
Cheek teeth prominently ridged.

Character suite A,

Neornithopods in which the antorbital vacuity is vestigial or absent.
Coronoid process a high prominence but not a projecting process. Epijugal
element present on jugal flange. Secondary palate extends posteriorly with
inclusion of the anterior ends of the maxillae, which are in contact (Galton 1973).
Vomer deep, vertically oriented sheet of bone bisecting anterior palatal vacuity.
Parietal and squamosal extensively produced posteriorly, overhanging occiput.
Scapula straight. Prepubis elongate. The 4th trochanter is not pendent.

While Steel (1969) commented on certain resemblances between pachy-
cephalosaurs and ceratopsians, until this relationship is more firmly established
and a monophyletic origin secured, this branch of the cladogram is unnamed.

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 291

Character suite B,

Infraorder PACHYCEPHALOSAURIA Maryanska & Osmolska, 1974 (nom. transl.
herein ex suborder Pachycephalosauria)

Moderately large neornithopods with the fronto-parietal region of the skull
greatly thickened and rugose, forming a dome-like prominence. Quadrate
markedly inclined, with narrow lateral temporal fenestra sloping forward to
beneath the orbit. Premaxilla reduced, maxilla extending up to meet the nasal.
Retroarticular process well developed. Ischium decurved.

Character suite C;

Family STENOPELIXIDAE Nopsca, 1917 (nom. correct. Kuhn, 1966)

Skull cap with two small domes, one on each frontal. Upper temporal
fenestra not reduced. Frontal not excluded from orbit margin by prefrontal. Pubis
reduced and excluded from acetabulum. Postpubis lacking.

The family Stenopelixidae is reintroduced because it is a senior synonym of
the Pachycephalosauridae. However, since the latter taxon 1s based largely upon
cranial material from the late Cretaceous whereas Stenopelix is a Weald genus, it
seems prudent to retain both families pending the discovery of additional
material.

Character suite D,

Family PACHYCEPHALOSAURIDAE Sternberg, 1945

Skull cap with single dome. Upper temporal fenestrae closed or vestigial.
Frontal excluded from orbit margin by fusion of prefrontal with supraorbital.

Character suite E

Infraorder CERATOPSIA Marsh, 1890 (nom. transl. herein
ex suborder Ceratopsia)

Upper jaw with rostral bone forming a prominent beak. Antorbital vacuities
vestigial or absent. Frontals and prefrontals enter orbital border. Retroarticular
process absent. Scapula of uniform width. Postpubis much reduced to obsolete.
Ischium decurved.

Character suite F

Superfamily PsITrACOSAUROIDEA new superfamily

Premaxillary teeth lacking. Jugal deep. Lateral temporal fenestra broad.
Manus with phalangeal formula 2—3—4—-1-0. Sacrum with 5-6 elements. Both
prepubis and postpubis short, slender. Ischium straight, long, with distal ends
blade-like and united in a broad symphysis. Crest-like greater trochanter. Four
distal tarsals.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family PsItrAcOSAURIDAE Osborn, 1923

Diagnosis as for superfamily.

Character suite G

Superfamily CERATOPSOIDEA Hay, 1902

Large-skulled quadrupeds with a cervical frill and various degrees of horn
formation. Maxillary and dentary teeth set in a groove. Atlas—axis complex,
together with 3rd and sometimes 4th cervical wholly or partially fused. Iliac
antitrochanter prominent.

Character suite H,

Family PROTOCERATOPSIDAE Granger & Gregory, 1923

Small ceratopsoids with at most an incipient horn core to the nasal.
Postorbitals arched and rugose but without horn development. Parietal-
squamosal frill short to very short. Coronoid process low. Sacrum with 6-8
elements. Ischium long, slender, almost straight.

Character suite I,

Family CERATOPSIDAE Marsh, 1888

Large to very large ceratopsoids with large external nares situated in well-
developed fossae. Cheek teeth double-rooted, up to 40 in each series. Premaxilla
edentulous. Nasals broad, usually with median horn core. Frontals and
prefrontals excluded from orbital border. Conspicuous posterior parietal-
squamosal frill. Postorbitals greatly expanded, with horns. Sacrum with 8-11
elements. Ischium short, broad, strongly decurved. Prepubis long. The
4th trochanter is reduced. Unguals hoof-shaped.

Discussion

On the basis of the primitive characters within the above recognized clades it
is possible to anticipate the ancestral ornithischian. It was a small biped with short
snout, small external nares, prominent antorbital vacuities, large rounded orbits
with sclerotic rings and each with a supraorbital. Dentition was heterodont and
bilaterally enamelled, with simple premaxillary teeth and up to 20 compressed,
single-rooted, spatulate cheek teeth. Wear surfaces were absent. The maxillary
teeth closed outside those of the dentary. The nasals were narrow and there was a
coronoid process. There were 24 presacrals (9 + 15), 4 sacrals and a long tail. The
scapula blade broadened posteriorly and there was an acromion process. Sternal
plates were present. There was a prominent deltopectoral crest to the humerus,
the latter longer than the radius and ulna. The manus was relatively small and
slender with digits IV and V lacking unguals, and a phalangeal formula of
2—3-—4—3-2. The unguals were claw-like. The ilium was low, with a long pointed

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 293

anterior process and a shorter, broader posterior process. It possessed a supra-
acetabular buttress but a brevis shelf was inconspicuous to absent. The pubis had
a short prepubis and a long postpubis. The femur was sinuous with a poorly
discriminated greater trochanter and a lesser trochanter adpressed to the shaft. It
lacked an anterior intercondylar groove. The tibia was stout and twisted, and the
hindlimbs were longer than the forelimbs. The astragalus and calcaneum were
separate and there were two distal tarsals. Metatarsals I-IV were slender and
elongate and the pedal phalangeal formula was 2—3—4—5-—0. Digit V was vestigial
and the pes was functionally tridactylous.

If the diagnostic ornithischian characters are removed from the above list,
the remaining features virtually diagnose the prosauropods and support Cooper’s
(1981) suggestion that the Ornithischia are a neotenous offshoot of prosauropod
stock. Moreover, it emphasizes the correctness of grouping the sauropodomorphs
and ornithischians together in the Ornithischiformes.

SYSTEMATIC DESCRIPTION
Order ORNITHOPODA Marsh, 1871
Suborder HyPSILOPHODONTIA new suborder
Superfamily HYPSILOPHODONTOIDEA new superfamily

Family Hypsilophodontidae Dollo, 1882

Discussion

When the early hypsilophodonts are stacked stratigraphically it is clear that
from an early stage there were a number of discrete phyletic lines. Thus, the late
Kimmeridgian Morrison Formation has yielded Othnielia, a typical hypsi-
lophodontid perhaps ancestral to Hypsilophodon itself, the primitive iguanodont
Camptosaurus, the fabrosaurid Nanosaurus and the aberrant hypsilophodontid
Dryosaurus. The coexistence of Othnielia and Dryosaurus points to an early
dichotomy of the Hypsilophodontidae and the writer thus proposes:

Subfamily Dryosaurinae new subfamily

Diagnosis

Moderately sized hypsilophodontids with edentulous premaxillae separating
maxillae and nasals. Ilium with very broad brevis shelf. Prepubis transversely
flattened. Ischium with proximally situated obturator process. Femur sigmoidal,
with weak to moderately developed anterior intercondylar groove, flat medial
surface to inner condyle, and with insertion area for m. caudifemoralis longus
well separated from 4th trochanter. Pes tridactylous, with metatarsals I and V
reduced to vestigial splints.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Were it not for the fact that-Dryosaurus occurs side-by-side with the
primitive iguanodont Camptosaurus it could be regarded as an ideal link between
hypsilophodonts and iguanodonts. The coexistence of these two taxa points to
convergence. Galton (1981) includes Dryosaurus within the Hypsilophodontidae,
Suggesting its derived characters are convergent towards the iguanodont
condition. On the basis of the available evidence, the writer would also assign
Valdosaurus and Kangnasaurus to this subfamily.

Genus Kangnasaurus Haughton, 1915
Type species Kangnasaurus coetzeei Haughton, 1915, by monotypy.

Kangnasaurus coetzeei Haughton, 1915
Figs 3-22
‘Kangnasaurus coetzeei Haughton, 1915: 19, figs 1-6. Steel, 1969: 19, fig. 8 (8-9).

Kangnasaurus . . . Lapparent & Lavocat (in Piveteau), 1955: 384. Romer, 1956: 629; 1966: 370.
Thulborn, 1974: 172. Taquet, 1975: 507, fig. 3.

Holotype
By original designation, the cheek tooth, SAM—2732 (Fig. 3).

Hypodigm

In addition to the holotype, the following material was collected from the
same well and in large part was used to supplement the original description of
IX, COEWCA:

SAM-2731 — right femur
2731a — proximal end of right femur
2731b — distal end of right femur
2731c — distal end of left femur
2731d — proximal end of right femur
2731e — articulated distal left femur and proximal portion of tibia
2731f — four articulated caudal vertebrae
2731g — distal end of left metatarsal

2731h — ?
27311 —?
2731) — distal portion of right tibia with articulated tarsus and proximal

portion of metatarsus

Locality

Rogers (1915) gave the locality as a well on the farm Kangnas, *. . . in a wide
shallow valley leading to the Orange River at Henkries’. The well from which
Kangnasaurus was obtained was sunk in the Koa River valley, about 7,5 km south
of Henkries Mond in Little Bushmanland, and 15 km south-west of Goodhouse

(Fig. 1).

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 295

Description
Tooth

A single cheek tooth is the only cranial fragment available (Fig. 3). It was
identified by Haughton (1915) as a right maxillary tooth but in hypsilophodontids
it is the dentary teeth that bear a strong median keel. The tooth is enamelled on
both surfaces, spatulate, and without a cingulum or constriction between root and
crown. The root seems to have been long and tapering, implying a fairly deep
dentary, with a subhexagonal cross-section and oval pulp cavity. The crown is
longitudinally curved and with a subrhomboidal medio-lateral profile. The convex
surface is strongly ridged, with a median keel forming a distinct spike to the
cutting edge, as in Hypsilophodon. In addition, there are strong ridges along both
the anterior and posterior margins, the stronger of which was believed by
Haughton (1915) to be the posterior one. Between these main ridges are a
number of slightly diverging subsidiary ridges, six on one side of the median keel
and eight on the other. The concave surface shows very faint and indistinct
longitudinal ridging and a pronounced biconcave wear facet indicating occlusion
with two teeth of the opposite jaw during mastication. Thus the maxillary and
dentary tooth rows were parasagittally offset, relative to one another. The cutting
edge of the crown is weakly serrated by the subsidiary ridges on the convex
surface.

Cervical vertebra

A single fragment of a cervical centrum (Fig. 4) gives the impression of
having been relatively long and strongly waisted at midlength, with a prominent
ventral keel. The articular face is convex, with a distinct pit just below centre. It
seems likely that this centrum was opisthocoelous, as in Camptosaurus and
Hypsilophodon.

Dorsal vertebra

A beautifully preserved dorsal centrum is platycoelous, almost amphipla-
tyan, with suboval anterior and posterior profiles (Fig. 5). Viewed laterally both
the anterior and posterior rims are broadly scarred for muscle attachment, and
there is a small nutritive foramen. Ventrally the centrum is narrowly rounded.
The ventral surface of the neural canal is pierced by a prominent, elongate
foramen, similar to but not as elongate as that of Dryosaurus lettowvorbecki
(Virchow) (Janensch 1955, fig. 22a).

Caudal vertebrae

There are six caudal vertebrae in the available material (Figs 6-8), four of
which comprise an articulated series. The largest is an isolated centrum (Fig. 6)
showing a subhexagonal cross-section at midlength, with the ventral surface
weakly grooved posteriorly. Three tiny foramina pierce the ventral surface and
there are distinct facets for chevron articulation posteriorly. In lateral view the

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Kangnasaurus coetzeei Haughton. The holotype, SAM—2732, an alleged maxillary tooth.
A. Labial view. B. Lingual view. C. Anterior view. Note the biconcave occlusal wear surface.
Bar scale = 5 mm.

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 297

Fig. 4. Kangnasaurus coetzeei Haughton. Fragment of a cervical vertebra.
A. Ventral view. B. Lateral view. C. Anterior view. Note the prominent ventral
keel. Bar scale = 25 mm.

anterior and posterior rims of the centrum are strongly scarred, while there is a
weak horizontal ridge at about midflank. The ventral surface of the neural canal is
perforated by two small foramina at about midlength. _

The articulated series (Fig. 7) lacks transverse processes and the neural
spines are very reduced. Since transverse processes are not lost before the twelfth
vertebra in Camptosaurus (Galton & Powell 1980) and Dryosaurus (Galton
1981), they are from the mid-portion of the series. The neurocentral sutures are
obsolete, their approximate positions marked by horizontal ridges. The lateral
surfaces of the centra are slightly concave while the ventral surface is weakly
grooved, giving the centra hexagonal cross-sections at midlength. There are
distinct chevron facets. The prezygapophyses are short, with subvertical articular
facets, and only just protrude beyond the anterior border of the centrum. The
postzygapophyses are naturally much longer, distinctly ridged and grooved
(Fig. 7), while there are two shallow dimples on the dorsal surface of the neural
arch, just behind the prezygapophyses, a feature also seen in the prosauropod
Massospondylus (Cooper 1981, fig. 15C).

Ribs
A proximal fragment of an anterior dorsal rib (Fig. 9) is the only bone
showing any features of note. The capitular pedicel and capitulum are broken off,

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig 5. Kangnasaurus coetzeei Haughton. A dorsal centrum. A. Ventral view. B. Lateral view.
C. Anterior view. D. Dorsal view. Bar scale = 25 mm.

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 299

Fig. 6. Kangnasaurus coetzeei Haughton. A mid-caudal centrum. A. Ventral view.
B. Lateral view. C. Dorsal view. D. Posterior view. Bar scale = 25 mm.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Kangnasaurus coetzeei Haughton. Arti-
culated mid-caudal vertebrae, SAM-—2731f.
A. Lateral view. B. Dorsal view. Note the
longitudinally grooved neural spines and sub-
pentagonal centra. Bar scale = 25 mm.

Fig. 8. Kangnasaurus coetzeei Haughton. A posterior caudal centrum. A. Lateral view.
B. Ventral view. C. Posterior view. Bar scale = 10 mm.

a

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 301

Fig. 9. Kangnasaurus coetzeei Haughton. Fragment of an anterior dorsal rib.
A. Anterior view. B. Ventral view. C. Posterior view. Bar scale = 25 mm.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

but there is a short tubercular pedicel with an elliptical tuberculum. Immediately
lateral to the tuberculum the rib has a V-shaped cross-section, with the apex
situated in the middle of the anterior surface. However, the anterior ridge quickly
shifts to an anteroventral position when the ventral surface of the rib becomes
gently concave.

Manus

A peculiar bean-shaped bone (Fig. 10) may be a manual phalanx or
metacarpal V. It is distinctly asymmetrical, dorsoventrally flattened, and with
rounded proximal and distal articular surfaces.

Pelvis

What may be a fragment of the proximal plate of an ischium is the only pelvic
remnant available. However, it is too scrappy and poorly preserved for proper
identification or description.

a

Fig. 10. Kangnasaurus coetzeei Haughton. Bone tentatively
identified as metacarpal V. A. Anconal view. B. Lateral view.
Bar scale = 10 mm.

Femur

The characters of the femur are well displayed by several specimens
(Figs 11-17). The femur is rather gracile (length/minimum transverse
width = 8,75), sigmodially curved in medial view (Fig. 12A) and strongly
expanded at both ends. The greater trochanter accounts for the proximal
parasagittal expansion and shows a gently curved dorsal surface in lateral view
(Fig. 12B). The lesser trochanter is closely adpressed to the greater trochanter, as
in Hypsilophodon, and not separated from it by a deep cleft as in Valdosaurus,
Dryosaurus, Camptosaurus, Callovosaurus and other ornithopods. There is some

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 303

Fig. 11. Kangnasaurus coetzeei Haughton.
A right femur, SAM~2731, anterior view.
Drawing reversed. Bar scale = 100 mm.

variation in the dorsal extent of the lesser trochanter. It is well below the level of
the greater trochanter in SAM-—2731 (Fig. 12), but only slightly so in SAM-—2731a
(Fig. 14). The 4th trochanter is not preserved in any of the available material,
although SAM-—2731 (Fig. 12) shows it to have been broad-based and with its
distal termination situated at midlength. It is likely to have been produced
posteriorly into a pendent, acuminate flange. The depression for the insertion of

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

mcl

Fig. 12. Kangnasaurus coetzeei Haughton. A right femur, SAM—2731.
A. Medial view. B. Lateral view. See also Fig. 11. Bar scale = 100 mm.

m. caudifemoralis longus is well separated from the 4th trochanter, in a
comparable position to Dryosaurus (cf. Galton 1981, fig. 14C, I). Distally there is
a moderately developed anterior intercondylar groove (Figs 13, 16), comparable
to the condition in some individuals of Dryosaurus (cf. Galton 1981, fig. 14J, L).
The distal end of the femur is strongly expanded transversely, with a very large
inner condyle and a smaller lateral condyle. As in Dryosaurus and Hypsi-
lophodon the medial surface of the inner condyle is flat. In the popliteal space

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 305

icg

Fig. 13. Kangnasaurus coetzeei Haughton. A. Distal end of a left femur,
SAM-2731c, anterior view. B. Anterior view of the proximal head of a
right femur, SAM-—2731a. Bar scale = 50 mm.

between the inner and outer condyles of SAM-—2731b (Fig. 15) is a rugose area
that marks the insertion of musculature referred to aponeurosis 1| in birds and the
prosauropod Massospondylus (Cooper 1981, fig. 84). Proximally the femoral
head is well developed, with a swollen rounded condyle, which is separated
dorsally from the greater trochanter by a pronounced groove (Fig. 17). The
posterior surface of the femoral condyle is distinctly concave, forming a
posteromedial lip.

- Tibia

A complete tibia is unknown, only the proximal and distal ends being
preserved. SAM—2731le comprises the distal end of a left femur articulated to the
proximal end of a tibia (Fig. 13A—B). The proximal head is strongly expanded,
with a well-developed cnemial crest that curves markedly outwards and is
separated from the accessory condyle by a broadly concave groove. Posteriorly,

the proximal articular surface shows well-developed inner and outer condyles, the
former the more prominent of the two. There is a pronounced accessory condyle

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. Kangnasaurus coetzeei Haughton. Proximal head of a right femur, SAM-—2731a.
A. Lateral view. B. Medial view. Note how the lesser trochanter does not reach the level of the
greater trochanter. See also Fig. 12B. Bar scale = 50 mm.

Fig. 15. Kangnasaurus coetzeei Haughton. Distal
end of a right femur, SAM-—2731b, in posterior
view. Bar scale = 50 mm.

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 307

Fig. 16. Kangnasaurus coetzeei Haughton. Distal
articular surface of a right femur, SAM-—2731b.
Bar scale = 50 mm.

Fig. 17. Kangnasaurus coetzeei Haughton. Proxi-

mal view of a right femur, SAM—273la. Note lack

of a broad cleft separating lesser and greater
trochanters. Bar scale = 50 mm.

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Kangnasaurus coetzeei Haughton. A-—B. Proxi-
mal articular surface and lateral view of the proximal head
of a right tibia, SAM-—273le. C. Distal end of a right tibia,
SAM-—2731j, in posterior view, with a cross-section of the
shaft. D. Distal articular surface of a right tibia,
SAM-2731j. E. Anconal and posterior views of distal
tarsal IV of the right pes, SAM—2731}. F. Palmar? view of
metatarsal V of the right pes of SAM-—2731). Bar scales in
5 mm divisions.

on the lateral surface of the outer condyle for articulation with the fibula, as in
Dryosaurus (Galton 1981, figs 16A, E-F, 19K) and Hypsilophodon (Galton
1974a, fig. 56A, E). Distally the preserved portion of SAM-—2731le tapers rapidly
and shows a suboval cross-section. SAM—2731j comprises the distal end of a right
tibia, together with loosely articulated tarsus and the proximal ends of the
metatarsals. The tibial fragment (Fig. 18C—D) is similar to both Hypsilophodon
(Galton 1974a, fig. 56) and Camptosaurus (Galton & Powell 1980, fig. 10L). It is
not significantly different from those tibiae of Dryosaurus in which a posterior
notch for the reception of the astragalus is lacking (Galton 1981). The anterior

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 309

surface of the outer malleolus is flat, for apposition with the distal end of the
fibula. The distal end of the tibia tapers proximally into a_ subtrigonal,
anteroposteriorly compressed shaft.

Astragalus

The astragalus is a thin cup of bone (Fig. 19D) which fitted snugly against the
distal end of the tibia. Its cupped dorsal surface is shallowly biconcave, with the
lateral cotylus the larger of the two. The distal roller is not completely exposed
but seems to have been smooth and weakly biconvex, with a shallow median
groove.

Fig. 19. Kangnasaurus coetzeei Haughton.

A-B. Right metatarsals III and IV of SAM-2731]j in
anconal and palmar views. C. Proximal profiles of
right metatarsals II-IV of SAM-2731j. E. Medial view
of proximal end of metatarsal II of SAM-2731).
Bar scales in 5 mm divisions.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Calcaneum

The calcaneum (Figs 20—21) is closely comparable to those of Hypsilophodon
(Galton 1974a, fig. S7A—E), Camptosaurus (Galton & Powell 1980, fig. 11F—G)
and Dryosaurus (Galion 1981, fig. 1ISA—C). The lateral surface is slightly
concave, almost flat, faintly scarred, and with a weak anteroventral rim which is
interrupted for a short distance ventrally. There is also a very weak dorsal rim,
just below the fibular facet. The most conspicuous feature of the medial surface is
a prominent tubercle, above which are four foramina. The distal roller is semi-
circular, smooth, and faintly corrugated, while the fibular facet is gently concave
and the tibial facet strongly concave.

Distal tarsals

A single distal tarsal (Fig. 18E) is preserved, which, to judge from its form
_ and position within the tarsus, is distal tarsal IV. Its medial edge is broken, but it
seems to have formed an elongate hemicylinder. The anterior surface bears two
deep grooves for ligament attachment, the dorsal surface is flat, the ventral
surface smoothly rounded for articulation with metatarsal IV, and the posterior
surface is shallowly concave.

Metatarsus

Apparently there were only four metatarsals to the pes of Kangnasaurus,
there being no evidence for metatarsal I; presumably it was obsolete. Metatarsals
II-IV are known only from their proximal ends. That of metatarsal II is not well
preserved but had a narrowly ovate proximal articular surface (Fig. 19C) whose
long axis seems to have been oriented subvertically. The proximal articular
surface of metatarsal III has an irregular profile, unlike that of any other
hypsilophodontid, while that of metatarsal IV is almost square but with a
depression on the lateral half of the surface for reception of distal tarsal IV. The
palmar surface of metatarsal IV is also deeply excavated to receive the vestigial
metatarsal V (Fig. 18F). The latter is a thin, narrow strap of bone, slightly curved
along its length, and similar in form and position to that of Dryosaurus (cf. Galton
(isi 1, sal),

Phalanges

The only pedal phalanx available is an asymmetrical, dorsoventrally
compressed ungual (Fig. 22) with deep medial and lateral grooves. The
asymmetry suggests it is from the right foot. It is non-diagnostic, but
indistinguishable from those of Dryosaurus and Camptosaurus.

Discussion

It is clear from the foregoing description of the known elements of
Kangnasaurus that it is a distinctive ornithopod, most closely allied to the
Dryosaurus—Hypsilophodon plexus. As such, its previous interpretation as an
iguanodontid (Lapparent & Lavocat in Piveteau 1955; Romer 1956, 1966; Steel

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON Sit|

Fig. 20. Kangnasaurus coetzeei Haughton. Left calceaneum. A. Lateral view. B. Medial view.
Bar scale = 10 mm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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A REVISION OF KANGNASAURUS COETZEEI HAUGHTON

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Fig. 22. Kangnasaurus coetzeei Haughton. Right pedal ungual. A. Lateral view.
B. Anconal view. Bar scale = 25 mm.

1969; Thulborn 1974; Taquet 1975) is incorrect (a conclusion independently
reached by P. M. Galton, in litt., 1981).

Kangnasaurus is readily distinguishable from [guanodon (Hooley 1925) by its
more gracile construction, sigmoidal femur with a shallow anterior intercondylar
groove, and markedly different cheek teeth. Callovosaurus is based upon a femur
(Galton 1980) whose greater trochanter does not show the parasagittal expansion
seen in Kangnasaurus, while its lesser trochanter is expanded transversely and
separated from the greater trochanter by a deep cleft.

The camptosaurid Muttaburrasaurus (Bartholomai & Molnar 1981) has a
much more robust femur than Kangnasaurus, with a distinctly smaller lesser
trochanter and a greater trochanter that does not show the parasagittal expansion
of Kangnasaurus. Its maxillary teeth are also rather different with up to 13 fine,
subparallel ridges on the labial surface, all of comparable strength.

Kangnasaurus differs from Camptosaurus (Galton & Powell 1980) in the
presence of an accessory (fibular) condyle to the tibia, a generally more sigmoidal
and less robust femur, and in having its lesser trochanter closely adpressed to the
greater trochanter.

Thescelosaurus (Parks 1926; Galton 19746; Morris 1976) is a hypsilophodon-
tid that has a much more robust femur than the South African genus, with the
4th trochanter extending on to a distal half of the shaft. In addition it has a
relatively well-developed first pedal digit and a peculiar ankle arrangement
(Morris 1976), in some species at least.

In its closely adpressed lesser and greater trochanters and flat medial surface
to the inner femoral condyle, Kangnasaurus is very similar to Hypsilophodon and
both show ‘high-spiked’ dentary teeth. Hypsilophodon, however, differs in
lacking an anterior intercondylar groove to the femur, the greater trochanter does

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

not show the degree of parasagittal expansion seen in Kangnasaurus, the cnemial
crest is not deflected outwards as strongly, metatarsal I is well developed, and the
dorsal centra of Hypsilophodon are amphicoelous whereas that referred to
Kangnasaurus is platycoelous.

Vectisaurus is a monotypic Wealden iguanodont recently redescribed by
Galton (1976). It differs from Kangnasaurus in lacking the slightly diverging
subsidiary ridges to the cheek teeth, and in having a deeply concave posterior
surface to its dorsal centra.

The hypsilophodontid Valdosaurus (Galton 1975) is based upon a femur that
differs from Kangnasaurus in having a very deep anterior intercondylar groove
and a deep cleft separating the lesser and greater trochanters. Othnielia is another
hypsilophodontid (Galton & Jensen 1973). It seems to have been a rather small
animal, distinguished from Kangnasaurus by a poorly developed anterior
_intercondylar groove to the femur, the transverse expansion of the lesser
trochanter, and the deep cleft separating lesser and greater trochanters.

Rozhdestvenskii (1966) described the iguanodontid Probactrosaurus from
the Lower Cretaceous of central Asia, a taxon widely regarded as close to the
ancestry of the hadrosaurs. Although much of the skeleton seems to have been
available, only the cranial characters were described in detail. However,
Probactrosaurus has a typical iguanodontid femur with very deep anterior
intercondylar groove. In addition, Rozhdestvenskii (1966) pointed out that the
remains described as the iguanodontid Sanpasaurus (Young 1944) are probably
those of a juvenile sauropod.

The highly derived camptosaurid Tenontosaurus (Ostrom 1970; Dodson
1980) is readily distinguished from Kangnasaurus by its massive femur with large
femoral head, the low position of the 4th trochanter, and the lack of an anterior
intercondylar groove.

Dysalotosaurus (Janensch 1955) has been shown to be a synonym of the
contemporaneous Morrison Dryosaurus (Galton 1977, 1981). The postcranial
osteology of both these taxa was described in detail by Galton (1981) and
Kangnasaurus is clearly a close ally. Femora of both taxa are sigmoidal, with a
moderately developed anterior intercondylar groove, a flat medial surface to the
inner condyle, and with the insertion area for m. caudifemoralis longus well
separated from the 4th trochanter. In addition, both Kangnasaurus and
Dryosaurus lack evidence of metatarsal I. However, the greater trochanter of
Dryosaurus does not show the parasagittal expansion seen in Kangnasaurus and is
separated from the lesser trochanter by a deep cleft in most individuals (Galton
1981). It is perhaps significant, however, that Shepard er al. (1977) figure a
Dryosaurus femur that seems to be comparable to that of Kangnasaurus.

Occurrence

Kangnasaurus coetzeei Haughton, the type and only species of the genus, is
known only from the northernmost Cape Province in South Africa. Its age is not

A REVISION OF KANGNASAURUS COETZEEI HAUGHTON 315

known but, based upon its stage of evolutionary development, an earliest
Cretaceous age seems reasonable.

SUMMARY

The osteology of Kangnasaurus coetzeei Haughton is described and figured in
detail. It is shown to be most closely allied to Dryosaurus from the late
Kimmeridgian of North America and Tanzania, and is thus transferred from the
Iguanodontidae to the Hypsilophodontidae. Together with Valdosaurus these
two genera are included in the new subfamily Dryosaurinae. Further work is
necessary to establish whether the Dryosaurinae are ancestral to later iguano-
donts or are merely a homoeomorphic hypsilophodontid development. Orni-
thischian taxonomy is currently unsatisfactory and character suites have been
used to construct a cladogram that is the basis for a new classification of
ornithischian dinosaurs.

ACKNOWLEDGEMENTS

Peter Galton obtained photographs of Kangnasaurus in 1976 with a view to
revising this material. However, since I had access to all the available material he
kindly agreed that I should work on it, for which I am grateful. In addition J
should like to thank Dr M. A. Cluver for allowing me to study the material,
Mr J. van den Heever for supervising the additional preparation and Mr F. Grine
for other assistance. Drs R. A. Thulborn and W. J. Morris are thanked for their
critical comments. The University of Durban-Westville is thanked for a grant in
aid of publication.

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A REVISION OF KANGNASAURUS COETZEEI HAUGHTON SH7i

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ZITTEL, K. A. von 1932. Text-book of palaeontology. Vol. 11. London: Macmillan & Co.

ABBREVIATIONS
apl — attachment area for aponeurosis |
Gi — chevron facet
fc — femoral condyle
ft — 4th trochanter
gt — greater trochanter
icg — intercondylar groove
Ic — lateral condyle
It — lesser trochanter
mc — medial condyle
mel — insertion area for m. caudifemoralis longus
ns — neural spine

prz — prezygapophysis

nh

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., Syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific namé must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata eons 1845: 37.
Leda plicifera A. Adams, : 50.
Laeda bicuspidata Hanley, 809: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new’species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text

>] e

eo 7) the Figure depicting C. namacolus ...’; ~. . . im C.\namacolus (Fig. 10)...

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

MICHAEL R. COOPER

A REVISION OF THE

ORNITHISCHIAN DINOSAUR
KANGNASAURUS COETZEEI HAUGHTON,
WITH A CLASSIFICATION OF

THE ORNITHISCHIA

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